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For Reference 


• "^'- I CALF 


1885- IQ56 


nfarr U'eUmnv, del. 


1 = Pholus pandorus. 

2 = Smerinthus geminatus. 

3 = Ainpelophaga versicolor. 
4=Marumba modesta. 

5 = Hemaris thysbe. 
6=Thyreus abbotti. 

American j^aturr Series; 

Group I. Classification of Nature 



Professor oj Entomology and Lecturer on Bionomics 
in Leliind Stanford Jr. University 





Copyright, 1904 






If man were not the dominant animal in the world, this would be the 
Age of Insects. Outnumbering in kinds the members of all other groups 
of animals combined, and showing a wealth of individuals and a degree 
of prolificness excelled only by the fishes among larger animals, and among 
smaller animals by the Protozoa, the insects have an indisputable claim on 
the attention of students of natural history by sheer force of numbers. But 
their claim to our interest rests on securer ground. Their immediate and 
important relation to man as enemies of his crops, and, as we have come to 
know only to-day, as it were, as a grim menace to his own health and life — 
this capacity of insects to destroy annually hundreds of millions of dollars' 
worth of grains and fruits and vegetables, and to be solely responsible for 
the dissemination of some of the most serious diseases that make man to 
suffer and die, forces our attention whether we will or not. Finally, the 
amazing variety and specialization of habit and appearance, the extraor- 
dinary adaptations and "shifts for a living" which insects show, make a 
claim on the attention of all who harbor the smallest trace of that "scientific 
curiosity" which leads men to observe and ponder the ways and seeming of 
Nature. Some of the most attractive and important problems which modern 
biological study is attacking, such as the significance of color and pattern, 
the reality of mechanism and automatism in the action and behavior of 
animals as contrasted with intelligent and discriminating performances 
the statistical and experimental study of variation and heredity, and other sub- 
jects of present-day biological investigation, are finding their most available 
material and data among the insects. 

This book is written in the endeavor to foster an interest in insect biology 
on the part of students of natural history, of nature observers, and of general 
readers; it provides in a single volume a general systematic account of all 
the principal groups of insects as they occur in America, together with special 
accounts of the structure, physiology, development and metamorphoses, and 
of certain particularly interesting and important ecological relations of insects 
with the world around them. Systematic entomology, economic entomology, 
and what may be called the bionomics of insects are the special subjects of 
the matter and illustration of the book. An effort has been made to put 
the matter at the easy command of the average intelligent reader; but it has 
been felt that a little demand on his attention will accomplish the result 
more satisfactorily than could be done with that utter freedom from etTort 

vi Prefatory Note 

with which some Nature-books try to disseminate knowledge. The few 
technical terms used are all explained in the text in connection with their 
first use, and besides are inserted in the Index with a specific reference, in 
black-faced type, to the explanation. So that the tyro reading casually in 
the book and meeting any of these terms apart from their explanation has 
only to refer to the Index for assistance. Readers more interested in accounts 
of the habits and kinds of insects than in their structure and physiology 
will be inclined lo skip the first three chapters, and may do so and still find 
the rest of the book "easy reading" and, it is hoped, not devoid of entertain- 
ment and advantage. But the reader is earnestly advised not to spare the 
little attention especially needed for understanding these first chapters, and 
thus to ensure for his later reading some of that quality which is among 
the most valued possessions of the best minds. 

In preparing such a book as this an author is under a host of obligations 
to previous writers and students which must perforce go unacknowledged. 
Some formal recognition, however, for aid and courtesies directly tendered 
by J. H. Comstock of Cornell University, whose entomological text-books 
have been for years the chief sources of knowledge of the insects of this 
country, I am able and glad to make. To my artist. Miss Mary Wellman, 
for her constant interest in a work that must often have been laborious and 
wearying, and for her persistently faithful endeavor toward accuracy, I extend 
sincere thanks. To Mrs. David Starr Jordan, who read all of the manuscript 
as a "general reader" critic, and to President Jordan for numerous sugges- 
tions I am particularly indebted. For special courtesies in the matter of 
illustrations (permission to have electrotypes made from original blocks) 
I am obhged to Prof. F. L. Washburn, State Entomologist of Minnesota (for 
nearly one hundred and fifty figures), Prof. M. V. Slingerland of Cornell 
University, Dr. E. P. Felt, State Entomologist of New York, Mr. Wm. 
Beutenmiiller, editor of the Journal of the New York Entomological Society, 
and Dr. Henry Skinner, editor of the Entomological News. 

Vernon L. Kellogg. 

Stanford University, California, 
June 1, 1904. 



I. The Structure and Special Physiology of Insects i 

II. Development and Metamorphosis of Insects 35 

III. Classification of Insects 52 

IV. The Simplest Insects (Order Aptera) 58 

V. May-flies (Order Ephemerida) and Stone-flies (Order Plecoptera). 65 

VI. Dragon-flies and Damsel-flies (Order Odonata) 75 

VII. The Termites or White Ants (Order Isoptera) 99 

VIII. Book-lice and Bark-lice (Order Corrodentia), and Biting Bird-lice 

(Order Mallophaga) iii 

IX. The Cockroaches, Crickets, Locusts, Grasshoppers, and Katydids 

(Order Orthoptera) 123 

X. The True Bugs, Cicadas, Aphids, Scale-insects, etc. (Order Hemip- 

tera), and the Thrips (Order Thysanoptera) 163 

XI. The Nerve-winged Insects (Order Neuroptera), Scorpion-flies 

(Order Mecoptera), and Caddis-flies (Order Trichoptera) 223 

XII. The Beetles (Order Coleoptera) 246 

XIII. The Two-winged Flies (Order Diptera) 301 

XIV. The Moths and Butterflies (Order Lepidoptera) 358 

XV. The Ichneumons, Gall-flies, Wasps, Bees, and Ants (Order Hymen- 

optera) 459 

XVI. Insects and Flowers 562 

XVII. Color and Pattern and their Uses 583 

XVIII. Insects and Disease 615 

Appendix: Collecting and Rearing Insects 635 

Index , 649 




ERHAPS no more uninteresting matter, for 
the general reader or entomological amateur, 
can be written about insects than a descrip- 
tive catalogue of the parts and pieces of the 
insect body. And such matter is practically 
useless because it doesn't stick in the reader's 
mind. If it is worth while knowing the 
intimate make-up of a house-fly's animated little body, it is worth 
getting this knowledge in the only way that will make it real, that is, 
by patient and eye-straining work with dissecting-needles and micro- 
scope. This book, anyway, is to try to convey some information about 
the kinds and ways of insects, and to stimulate interest in insect life, rather 
than to be a treatise on insect organs and their particular functions. Life 
is, to be sure, only the sum of the organic functions, but this sum or com- 
bination has an interest disproportionate to that of any of its component 
parts, and has an aspect and character which cannot be foretold in any com- 
pleteness from ever so careful a disjoined study of the particular functions. 
And so with the body, the sum of the organs: it is the manner and seeming 
of the body as a whole, its symmetry and exquisite adaptation to the special 
habit of life, the fine delicacy of its colors and pattern, or, at the other 
extreme, their amazing contrasts and hizarrerie, on which depend our first 
interest in the insect body. A second interest, although to the collector and 
amateur perhaps the dominant one, comes from that recognition of the 
differences and resemblances among the various insects which is simply 
the appreciation of kinds, i.e., of species. This interest expanded by oppor- 
tunity and observation and controlled by reason and the habit of order and 
arrangement is, when extreme, that ardent and much misunderstood and 
scoffed at but ever-impelling mainspring of the collector and classifier. 

2 The Structure and Special Physiology of Insects 

Of all entomologists, students of insects, the very large majority are col- 
lectors and classifiers, and of amateurs apart from the few who have "crawl- 
eries" and aquaria for keeping alive and rearing " worms" and water-bugs 
and the few bee-keepers who are more interested in bees than honey, prac- 
tically all are collectors and arrangers. So, as collecting depends on a 
knowledge of the life of the insect as a whole, and classifying (apart from 
certain primary distinctions) on only the external structural character of 
the body, any detailed disquisition on the intimate character of the insec- 
tean insides would certainly not be welcome to most of the users of this 

That insects agree among themselves in some important characteristics 
and differ from all other animals in the possession of these characteristics 
is implied in the segregation of insects into a single great class of animals- 
Class here is used with the technical meaning of the systematic zoologist- 
He says that the animal kingdom is separable into, or, better, is composed 
of several primary groups of animals, the members of each group possessing 
in common certain important and fundamental characteristics of structure 
and function which are lacking, at any rate in similar combination, in all 
other animals. These primary groups are called phyla or branches- All 
the minute one-celled animals, for example, compose the phylum Protozoa 
(the simplest animals); all the starfishes, sea-urchins, sea-cucumbers, and 
feather-stars, which have the body built on a radiate plan and have no back- 
bone, and have and do not have certain various other important things, 
compose the phylum or branch Echinodermata; all the back-boned ani- 
mals and some few others with a cartilaginous rod instead of a bony column 
along the back compose the class Chordata; all the animals which have 
the body composed of a series of successive rings or segments, and have 
pairs of jointed appendages used as feet, mouth-parts, feelers, etc., aris- 
ing from these segments, compose the phylum Arthropoda. There are 
still other phyla — but I am not writing a zoology. The insects are Arthro- 
poda; and any one may readily see — it is most plainly seen in such forms as 
a locust, or dragon-fly, or butterfly, and less plainly in the concentrated 
knobby little body of a house-fly or bee — that an insect's body shows the 
characteristic arthropod structure; it is made up of rings or segments, and 
the appendages, legs for easiest example, are jointed. An earthworm's 
body is made up of rings, but it has no jointed appendages. A worm is 
therefore not an arthropod. A crayfish, however, is made up of distinct 
successive body-rings, and its legs and other appendages are jointed. And 
so with crabs and lobsters and shrimps. And the same is true of thousand- 
legged worms and centipeds and scorpions and spiders. All these creatures, 
then, are Arthropods. But they are not insects. So all the back-boned 
animals, fishes, amphibians, reptiles, birds, and mammals are Chordates, 

The Structure and Special Physiology of Insects 3 

but they are not all birds. The phylum Chordata is subdivided into or 
composed of the various classes Pisces (fishes), Aves (birds), etc. And 
similarly the phylum Arthropoda is composed of several distinct classes, 
viz.: the Crustacea, including the crayfishes, crabs, shrimps, lobsters, 
water-fleas, and barnacles; the Onychophora, containing a single genus 
(Peripatus) of worm-like creatures; the Myriapoda, including the thousand- 
legged worms and centipeds; the Arachnida, including the scorpions, spiders, 
mites, and ticks; and finally the class Insecta (or Hexapoda, as it is some- 
times called), whose members are distinguished from the other Arthro- 




tarsal segments 

Fig. I. — Locust (enlarged) with external parts named. 

pods by having the body-rings or segments grouped into three regions, called 
head, thorax, and abdomen, by having jointed appendages only on the body- 
rings composing the head and thorax (one or two pairs of appendages may 
occur on the terminal segments of the abdomen), and by breathing by means 
of air-tubes (tracheje) which ramify the whole interior of the body and 
open on its surface through paired openings (spiracles). The insects also 
have three pairs of legs, never more, and less only in cases of degeneration, 
and by this obvious character can be readily distinguished from the Myria- 
pods, which have many pairs, and the Arachnids, which have four pairs. 
Centipeds are not insects, nor are spiders and mites and ticks. What 
are insects most of this book is given to showing. 

To proceed to the classifying of insects into orders and families and 
genera and species inside of the all-including class is the next work of the 
collector and classifier. And for this— if for no other reason— some further 
knowledge of insect structure is indispensable. The classification rests 

4 The Structure and Special Physiology of Insects 

mostly on resemblances and differences in corresponding parts of the body, 
apparent in the various insect kinds. What these parts are, with their names 
and general characters, and what their particular use and significance are, 
may be got partly from the following brief general account, and partly from 
the special accounts given in connection with special groups of insects else- 
where in this book. A little patience and concentration of attention in 
the reading of the next few pages will make the reader's attention to the 
rest of the book much simpler, and his understanding of it much more 

The outer layer of the skin or body-wall of an insect is called the cuticle, 
and in most insects the cuticle of most of the body is firm and horny in char- 

FiG. 2. — Longitudinal section of anterior half of an insect, Menopon titan, to show chitin- 
ized exoskeleton, with muscles attached to the inner surface. (Much enlarged.) 

acter, due to the deposition in it, by the cells of the skin, of a substance called 
chitin. This firm external chitinized * cuticle (Fig. 2) forms an enclosing 
exoskeleton which serves at once to protect the inner soft parts from injury 

Fig. 3.- 

-Bit of body-wall, greatly magnified, of larva of blow-fly, Calliphora erythrocephala, 
to show attachment of muscles to inner surface. 

and to afford rigid points of attachment (Figs. 2, 3 and 4) for the many small 
but strong muscles which compose the insect's complex muscular system. 
Insects have no internal skeleton, although in many cases small processes 
project internally from the exoskeleton, particularly in the thorax or part 

* It is not certainly known whether the cuticle is wholly secreted by the skin cells, or 
is in part composed of the modified external ends of the cells themselves. 

The Structure and Special Physiology of Insects 5 

of the body bearing the wings and legs. Where the cuticle is not strongly 

chitinized it is flexible (Fig. 6), thus permitting 

the necessary movement or play of the rings 

of the body, the segments of the legs, antennae 

and mouth-parts, and other parts. The small 

portions of chitinized cuticle thus isolated or 

made separate by the thin interspaces or sutures 

4. ■ Fig. 5. 

jri(;_ ^_ — Diagram of cross-section through the thorax of an insect to show leg and wing 

muscles and their attachment to body-wall, h., heart; al.c, alimentary canal; v.n.c. 

ventral nerve-cord; w., wing; /., leg; w., muscles. (Much enlarged; after Graber.) 
PiQ J — Left middle leg of cockroach with exoskeleton partly removed, showing muscles. 

(Much enlarged; after Miall and Denny.) 

are called sclerites, and many of them have received specific names, while 
their varying shape and character are made use of in distinguishing and 
classifying insects. 

Fig. 6. — Chitinized cuticle from dorsal wall of two body segments of an insect, showing 
sutures (the bent places) between segmental sclerites. Note that the cuticle is not 
less thick in the sutures than in the sclerites, but is less strongly chitinized (indi- 
cated by its paler color). 

The whole body is composed fundamentally of successive segments 
(Figs. I and 7), which may be pretty distinct and similar, as in a caterpillar 
or termite or locust, or fused together, and strongly modified, and hence 
dissimilar, as in a house-fly or honey-bee. The segments, originally five 
or six, composing the head, are in all insects wholly fused to form a single 
box-like cranium, while the three segments which compose the thorax are 
in most forms so fused and modified as to be only with difficulty distinguished 
as originally independent body-rings. On the other hand, in most insects 

6 The Structure and Special Physiology of Insects 

the segments of the abdomen retain their independence and are more or 

compound eye, 
prothorax^ ' 



tarsal segments 

Fig. 7. — Body of the monarch butterfly, Anosia plexippiis, with scales removed to show 
external parts. (Much enlarged.) 

less similar, thus preserving a generalized or ancestral condition. On the 
head are usually four pairs of jointed appendages (Fig. 8), viz., the 

antennae and three pairs of mouth-parts, 
known as mandibles, maxillae, and labium or 
under-lip. Of these the mandibles in most 
cases are only one-segmented, while the two 
members of the labial pair have fused along 
their inner edges to form the single lip-like 
labium. The so-called upper lip or labrum, 
closing the mouth above, is simply a fold of 
the skin, and is not homologous, as a true 
appendage or pair of appendages, with the 
other mouth-parts. In some insects with highly 
modified mouth structure certain of the parts 
may be wholly lost, as is true of the mandibles 
in the case of all the butterflies. The head 
bears also the large compound eyes and * the 
smaller simple eyes or ocelli (for an account of 
the eyes see p. 30). Attached to the thorax are 
three pairs of legs, which are jointed appendages, 
homologous in origin and fundamental struc- 

FiG. 8. — Dorsal aspect of head 
of dobson-fly, Corydalis cor- 
nuta, female, showing mouth- 
parts. Ih., labrum, removed; 
md., mandible; mx., maxilla; 
li., labium; gl, glossae of la- 
bium; St., stipes of maxilla; 
mxp., palpus of maxilla; ant., 

ture with the mouth-parts and antennas, and two pairs of wings (one or 

The Structure and Special Physiology of Insects 7 

both pairs may be wanting) which are expansions of the dorso-lateral 
skin or body-wall, and are not homologous with the jomted ventra 
appendages. The thorax usually has its first or most anterior segmentl 
the prothorax, distinct from the other two and freely movable, while 
the hinder two, called meso- and meta-thoracic segments, are usually, 
enlarged and firmly fused to form a box for holding and giving attachment 
to the numerous strong muscles which move the wings and legs. The 
abdomen usually includes ten or eleven segments without appendages or 
projecting processes except in the case of the last two or three, which bear 
in the female the parts composing the egg-laying organ or ovipositor, or 

Fig. 9. Fig. lo. 

Fig. 9.— Head, much enlarged, of mosquito, Culex sp., showing piercing and sucking 
mouth-parts. (After Jordan and Kellogg.) , tvt . .u u . * „,»i 

Fig 10— Head and mouth-parts of honey-bee, much enlarged. Note the short, trowel- 
" like mandibles for moulding wax when building comb, and the extended proboscis 
for sucking flower-nectar. (Much enlarged.) 

in certain insects the sting, and in the male the parts called claspers, cerci, 
etc., which are used in mating. On the abdomen are usually specially notice- 
able, as minute paired openings on the lateral aspects of the segments, the 
breathing-pores or spiracles, which admit air into the elaborate system of 
tracheae or air-tubes, which ramify the whole internal body (see p. 19). 

Of all these external parts two groups are particularly used in schemes 
of classification because of their structural and physiological importance 
in connection with the special habits and functions of insect life, and because 

8 The Structure and Special Physiology of Insects 

of the pronounced modifications and differences in their condition: these 
are the mouth-parts and the wings. 

Insects exhibit an amazing variety in food-habit: the female mosquito Hkes 
blood, the honey-bee and butterfly drink flower-nectar, the chinch-bug sucks 
the sap from corn-leaves, the elm-leaf beetle and maple worm bite and chew 
the leaves of our finest shade-trees, the carrion-beetles devour decaying 
animal matter, the house-fly laps up sirup or rasps off and dissolves loaf- 
sugar, the nut- and grain-weevils nibble the 
dry starchy food of these seeds, while the 
apple-tree borer and timber-beetles find 
sustenance in the dry wood of the tree- 
trunks. The biting bird-lice are content 
with bits of hair and feathers, the clothes- 
moths and carpet-beetles feast on our :ugs 
and woolens, while the cigarette-beetle has 
the depraved taste of our modern vouth. 

Fig. II. 

Fig. II. — Mouth-parts, much enlarged, of the house-fly, Musca domestica. mx.p., maxil- 
lary palpi; ih., labrum; li., labium; la., labellum. 

Fig. 12. — Head and mouth-parts, much enlarged, of thrips. ant., antenna; lb., labrum; 
md., mandible; mx., maxilla; mx.p., maxillary palpus; li.p., labial palpus; m.s], 
mouth-stylet. (After Uzel; much enlarged.) 

With all this variety of food, it is obvious that the food-taking parts must 
show many differences; one insect needs strong biting jaws (Fig. 8), another 
a sharp piercing beak (Figs. 9, 13, and 14), another a long flexible sucking 
proboscis (Figs. 10 and 16), and another a broad lapping tongue (Fig. n). 
Just this variety of structure actually exists, and in it the classific entomolo- 
gist has found a basis for much of his modern classification. 

Throughout all this range of mouth structure the insect morphologists 
and students of homology, beginning with Savigny in 18 16, have been able 
to trace the fundamental three pairs of oral jointed appendages, the mandi- 
bles, maxillae, and labium. Each pair appears in widely differing condi- 
tions; the mandibles may be large strong jaws for biting and crushing, as 
with the locust, or trowel-like, for moulding wax, as with the honey-bee, or 

The Structure and Special Physiology of Insects 9 

long, flat, slender, and saw-toothed, as with the scorpion-flies, or needle-like, 
as in all the sucking bugs, or reduced to mere rudiments or wholly lacking, 
as in the moths and butterflies. Similarly with the other parts. But by 
careful study of the comparative anatomy of the mouth structure, and par- 
ticularly by tracing its development in typical species representing the 
various types of biting, sucking, and lapping mouths, all the various kinds of 
mouth structure can be compared and the homologies or structural cor- 
respondences of the component parts determined. Figs. 8 to 16 illustrate 


Fig. 13. Fig. 14. 

Fig. 13. — Seventeen-year cicada, Cicada septcndecim, sucking sajj from twig. (After 

Quaintance; natural size.) 
Fig. 14. — Section of twig of Carolina poplar showing beak of cicada in position when 

sucking. (After Quaintance; much enlarged.) 
Fig. 15. — Mouth-parts, much enlarged, of net-winged midge, Bibicocephala doanei, 

female, md., mandible; mx., maxilla; tnx.L, maxillary lobe; mx.p., maxillary 

palpus; //., labium; hyp., hypopharynx; pg., paraglossa of labium; l.ep., labrum 

and epipharynx. 

examples of different mouth structures, with the corresponding parts similarly 

The most conspicuous structural characteristic of insects is their poses- 
sion of wings. And the wings undoubtedly account for much of the success 
of the insect type. Insects are the dominant animal group of this age, as 
far as number of species constitutes dominance, their total largely sur- 
passing that of the species of all the other kinds of living animals. Flight 
is an extremely effective mode of locomotion, being swift, unimpeded by 
obstacles, and hence direct and distance-saving, and an animal in flight 
is safe from most of its enemies. The wings of insects are not modified true 
appendages of the body, but arise as simple sac-like expansions (Fig. 17) 
of the body-wall or skin much flattened and supported by a framework of 

lo The Structure and Special Physiology of Insects 


strongly chitinized lines called veins. These veins are corresponding cutic- 

ular thickenings, in the upper and lower walls of 
the flattened wing-sac, which protect, while the 
wing is forming, certain main tracheal trunks that 
carry air to the wing-tissue. After the wing is 
expanded and dry, the tracheae mostly die out, and 
the veins are left as firm thick-walled branching 
tubes which serve admirably as a skeleton or 
framework for the thin membranous wings. It 
has been found that despite the obvious great 
variety in the venation, or number and arrange- 
ment of these veins of the wing, a general type- 
plan of venation is apparent throughout the insect 
class. The more important and constant veins have 
been given names, and their branches numbers 
(Fig. i8). By the use of the same name or 
number for the corresponding vein throughout all 
the insect orders, the homologies or morphological 
correspondences of the veins as they appear in the 
variously modified wings of the different insects 
are made apparent. Many figures scattered through 
this book show the venation of insects of 
different orders, and the corresponding 
lettering and numbering indicate the 
homologies of the veins. As the wing 
venation presents differing conditions 
readily noted and described, much use is 
made of it in classification. 

The differences in the wings them- 
selves, that is, in number, relative size 
of fore and hind wings, and in struc- 
ture, i.e., whether membranous and 
delicate, or horny and firm, etc., have 

, , . , . always been used to distinguish the 

Fig. i6. — Sphinx moth, showing proboscis; -^ ^ r ■ 

at left the proboscis is shown coiled up larger groups, as orders, of msects, 
on the under side of the head, the nor- ^nd the first classification, that of 
mal position when not in use. (Large _ . , \ j- -j i.i. i 

figure, one-half natural size; small fig- Lmnsus (1750 app.), divides the class 
ure, natural size.) into orders almost solely on a basis 

of wing characters. The ordinal names expressed, to some degree, the 
differences, as Diptera,* two-winged; Lepidoptera, scale-winged; Coleoptera, 
sheath-winged, and so on. As a matter of fact, there may be much differ- 
* The derivation of the Lianaean ordinal names is given on p, 223. 

Fig. 16. 

The Structure and Special Physiology of Insects 1 1 

ence in the wings within a single order; most beetles, for example, have 
four wings, but some have two and some none. There are indeed wingless 
species in almost every insect order. But a typical beetle has quite dis- 
tinctive and commonly recognized wing characters; that is, it has two pairs 
of wings, the fore pair being greatly thickened, and developed to serve as 
sheaths for the larger, membranous under-pair, which are the true flight 
wings. Similarly, practically all moths and butterflies have two pairs of 

Ik \ 

Fig. 17. Fig. li 

Fig. 17. — Wing of cabbage-butterfly, Pieris rapa, in early sac-like stage, tr., trachea; 

//., tracheoles; l.v., lines of future veins. (After Mercer; greatly magnified.) 
Fig. 18. — Diagram of wings of monarch butterfly, Anosia plexippits, showing venation. 

c, costal vein; s.c, subcostal vein; r., radial vein; cii., cubital vein; a., anal veins. 

In addition, most insects have a vein lying between the subcostal and radial veins, 

called the median vein. (Natural size.) 

membranous wings completely covered above and below by small scales, 
which give them their distinctive color and pattern. 

The exoskeleton, or cuticle, of the insect body is sometimes nearly 
smooth and naked, but usually it is sculptured by grooves and ridges, punc- 
tures or projections, and clothed with hairs or those modified flattened hairs 
known as scales (especially characteristic of butterflies and moths). This 
clothing of hairs or scales, or the skin itself, is variously colored and pat- 
terned, often with the obvious use of producing protective resemblance or 
mimicry, but often without apparent significance. (For an account of the colors 
and patterns of insects and their uses see Chapter XVII.) The hairs may serve 
for protection, or may be tactile organs, or even organs of hearing (see p. 26). 
The projecting processes may be spines or thorns or curious and inexplicable 

I 2 The Structure and Special Physiology of Insects 

knobs and horns. The rhinoceros-beetle (Dynastes) (Fig. 19) and the sacred 
scarabeus are famihar examples of insects with such prominent processes. 

The insect body, as a whole, appears in great variety of form and range 
of size, as our knowledge of the variety of habit and habitat of insects would 
lead us to expect. In size they vary from the tiny four-winged chalcids 
which emerge, after their parasitic immature life, from the eggs of other 
.nsects, and measure less than a millimeter in length, to the giant Phasmids 


Fig. 19. — Rhinoceros-beetle, Dynastes tityriis, showing chitinous horns. 

(walking-sticks) of the tropics, with their ten or twelve inches of body length, 
and the great Formosan dragon-flies with an expanse of wing of ten 
inches. A Carboniferous insect like a dragon-fly, known from fossils found 
at Commentry, France, had a wing expanse of more than two feet. 
Insects show a plasticity as to general body shape and appearance that results 
in extreme modifications corresponding with the extremely various habits 
of life that obtain in the class. Compare the delicate fragility of the gauzy- 
winged May-fly with the rigid e.xoskeleton and horny wings of the water- 
beetle; the long- winged, slender-bodied flying-machine we call a dragon- 
fly with the shovel-footed, half-blind, burrowing mole-cricket; the plump, 
toothsome white ant that defends itself by simple prolificness with the spare, 
angular, twig-like body of the walking-stick with its effective protective 
resemblance to the dry branches among which it lives. Compare the leg- 
less, eyeless, antennaless, wingless, sac-like degraded body of the orange- 
scale with the marvelous specialization of structure of that compact expo- 
nent of the strenuous insect life, the honey-bee; contrast the dull colors of the 
lowly tumble-bug with the flashing radiance of the painted lady-butterfly. 
But through all this variety of shape and pattern, complexity and degenera- 
tion, one can see the simple fundamental insect body-plan; the successive 
segments, their grouping into three body-regions, the presence of segmented 
appendages on head and thorax and their absence on abdomen (e.xcept 
perhaps in the terminal segments), and the modification of these append- 
ages into antennae and mouth-parts on the head, legs on the thorax, and 
ovipositor, sting, or claspers in the abdomen. 

In the character of the structure and functions of the internal organs 

The Structure and Special Physiology of Insects i 3 

or systems of organs of insects, a special interest attaches to the conditions 
shown by the circulatory and respiratory systems, and by the special sense- 


Fig. 20. — Diagram of lateral interior view of monarch butterfly, Anosia plexippus, show- 
ing the internal organs in their natural arrangement, after the removal of the right 
half of the body-wall together with the tracheje and fat body; I to III, segments 
of the thorax; i to g, segments of the abdomen. Alimentary Canal and Appen- 
dages: ph., pharynx; sd. and sgl., salivary duct and gland of the right side; oe., 
oesophagus; f.r., food -reservoir; St., stomach; i., small intestine; c, colon; r., rec- 
tum; a., anus; m.v., Malpighian tube. Haemal System: h., heart or dorsal vessel; 
ao., aorta; a.c, aortal chamber; Nervous System (dotted in figure): br., brain; 
g., subcesophageal ganglion; l.g., compound thoracic ganglia; fl,g.,, ag.^, first and 
fourth abdominal ganglia. Female Reproductive Organs: cp., copulatory pouch; 
v., vagina; o., oviduct, and oo., its external opening; r.ov., base of the right ovarian 
tubes turned down to expose the underlying organs; l.ov., left ovarian tubes in posi- 
tion, and ov.c, their termination and four cords; sp., spermatheca;^, part 
of the single accessory gland;^, one of the paired accessory glands; only the 
base of its mate is shown. Head: a., antenna; mx., proboscis, p., labial palpus. 
(After Burgess; three times natural size.) 

organs and their manner of functioning. The muscular system varies from the 
.simple worm-like arrangement of segmentally disposed longitudinal and 
ring muscles possessed by the caterpillars, grubs, and other worm-like larvae, 
to the complicated system of such 
specialized and active forms as the 
honey-bee and house-fly. Lyonnet 
describes about two thousand dis- 
tinct muscles in the caterpillar 'of 
the goat-moth. Insect muscles are 
similar, in their finer structure, to 
those of other animals, most of Fig. 21. — Bit of muscle of a biting bird-louse, 
them being composed of finely Eurymetopus tatcrus. (Greatly magnified.) 
cross-striated fibers (Figs. 21 and 22) held together in larger or smaller 
masses and attaching to the rugosities of the inner surface of the exo- 
skeleton. The muscle substance, when fresh, is peculiarly transparent 
and delicate-looking, but it has great contractile power. 

The alimentary canal (Figs. 23-27), like that of other animals, is a tube 
but little longer than the body in flesh-eating forms, and much longer in 
plant-feeders; it runs, more or less curving and coiled, through the body 
from mouth to anal opening, which lies in the last segment of the abdomen. 

14 The Structure and Special Physiology of Insects 
This tube is expanded variously to form crop, gizzard, or stomach, and 

Fig. 22. — Diagrammatic figures of bits of insect muscle, variously treated. 
Gehuchten; greatly magnified.) 

(After Van 

Fig. 23. — Alimentary 
canal of a locust. At 
upper end the oesoph- 
agus, then the ex- 
panded crop, then sev- 
eral large gastric coeca, 
then the true stomach, 
the thread-like Malpig- 
hian tubules, the bent 
intestine, and the ex- 
panded rectum. (After 
Snodgrass; enlarged.) 

contracted elsewhere to be oesophagus or intestine. 
One or two pairs of saHvary glands pour their fluid into 
the mouth, while the digesting stomach or ventriculus 
usually possesses two or more pairs of diverticula known 
as gastric coeca, which are lined with glands believed 
to secrete special digestive fluids. Neither liver 
nor kidneys are present in the insect body, but the 
secretory function of the latter are undertaken by a 
number of usually long thread-like tubular diverticula 
of the intestine known as Malpighian tubules. The 
intestine itself is usually obviously made up of three 
successive parts, a large intestine, small intestine, 
and rectum. There are also 
present not infrequently in- 
testinal ca?ca. 

Two striking peculiarities 
about the reproductive system 
of insects are the possession 
by the female of one or more 
spermathecce (Fig. 66, r.s.) in 
which the male fertilizing 
cells, the spermatozoa, are re- 
ceived and held, and the com- 
pletion of all the envelopes of „ t^- .. r 
^ .... , Fig. 24. — Dissection of 
the egg, mcludmg the outer cockroach to show {al.c.) 
hard shell, before its specific alimentary canal. (After 
. ... . , , T^ Hatschek and Cori: twice 
fertilization takes place, rer- natural si^e.) 

The Structure and Special Physiology of Insects i 5 

tilization is itself accomplished in the lower end of the egg-duct just before the 
egg is laid, by the escape of spermatozoa from the spermatheca (the female 




Fig. 25. Fig. 26. 

Fig. 25. — Alimentary canal of larva of harlequin-fly (Chironovius sp.). oes., oesophagus; 
s.g., salivary gland; ca., cardiac chamber of stomach; mt., Malpighian tubules; ch., 
intestinal chamber; si., small intestine; col., colon. (After Miall and Hammond; 
much enlarged.) 

Fig. 26. — AUmentary canal of two species of thrips; at left Trichothrips copiosa, male, 
at right Aelothrips fasciata. sal.g., saHvary gland; oes., oesophagus; prov., proven- 
triculus; vent., ventriculus; m.t., Malpighian tubules; int., intestine; rec, rectum. 
(After Uzel; greatly enlarged.) 

having of course previously mated) and their entrance into the egg through a 
tiny opening, the micropyle (Fig. 67), in the egg-shell and inner envelopes. 
A queen bee mates but once, but she may live for four or five years after 
this and continue to lay fertilized eggs during all this time. She must 

1 6 The Structure and Special Physiology of Insects 

receive several million spermatozoa at mating, and retain them alive in the 
spermatheca during these after-years. 


FiG. 27. — Alimentary canal of dobson-fly, Cor)'(fa/wconi?</a. A, larva; B, adult; C, pupa; 
oes., oesophagus; prov., proventriculus; g.c, gastric coeca; vent., ventriculus; r.g., 
reproductive gland; vi.t., Malpighian tubules; int., intestine; iut.c, intestinal 
coecum; rec, rectum; drg., oviduct. (After Leidy; twice natural size.) 

The circulatory system of insects presents two particular features of inter- 
est in that the blood does not, as in our bodies, carry oxygen to the tissues, and 

Fig. 28. — Cross-section and longitudinal section of salivary gland of giant crane-fly, 
Holorusia rubiginosa. (Greatly magnified.) 

that there is a contractile pulsating heart-like organ, but no arteries or veins. 
The so-called heart is a delicate-walled, narrow, subcylindrical vessel com- 
posed of a series of most commonly from three to eight successive cham- 
bers lying longitudinally along the median line just underneath the dorsal 
wall of the abdomen and thorax (Figs. 30 and 31). Each chamber opens, 
guarded by a simple valvular arrangement (Fig. 33), into the chambers 

The Structure and Special Physiology of Insects 17 
behind and before it, the posterior one being closed behind and the anterior 

Fig. 29. — Cells of digestive epithelium of stomach (ventriculus) of crane-fly, Ptychoptera 
sp., showing secretion of digestive fluids, or expulsion of cell-content. (After Van 
Gehuchten; greatly magnified.) 

one extending forward into or near the head as a narrowed tubular anterior 
portion, which is sometimes called the 
aorta. From the anterior open end of 
this aorta the blood, forced by pulsations 
of the heart-chambers, which proceed 
rhythmically from the posterior one 
forward, pours out into the body-cavity, 
proceeding in more or less regular cur- 
rents or paths, but never enclosed in 
arterial vessels, bathing all the tissues, 
and carrying food to them. Finally 
taking up fresh supplies of food by bath- 
ing the food-absorbing walls of the 
alimentary canal, it enters the chambers 
of the heart through lateral openings in 
these (either at the middle or anterior end 
of each), which thus estabhsh communi- 
cation between the body-cavity and heart- Fig. 30. Fig. 31. 
The blood receives no more oxygen than Fig. 30.— Diagram of circulatory 
it needs for its own use, and thus does ^^^gX t mrchimS'dis'al 
not play nearly so complex a function in vessel, or heart, with single artery. 
the insect's body as in ours. And this Ar^ws^mdicate^direc^on „, blood- 

simplicity of function probably explains pj^ 31.— Dissection showing dorsal 

in some degree the extreme primitiveness vessel, or heart, of locust, Dis- 
,^ . , . ,^ , sostezra Carolina. (After Snodgrass; 

of the make-up of the circulatory system. ^^j^^ natural size.) 

It will be seen that the respiratory 

system, on the other hand, is particularly highly developed, as it devolves 

I 8 The Structure and Special Physiology of Insects 

Fig. 32. 

Fig. 33. 

Fig. 32. — Portion of dorsal vessel and pericardial membrane of locust, Dissosteira caro* 

Una. (After Snodgrass; greatly magnified.) 
Fig. 33. — Cross-section of dorsal vessel or heart in pupa of tussock-moth, Hemerocampa 

leiicostigma, showing valves. (Greatly magnified.) 


Fig. 34. 

Fig. 35. 

Fig. 36. 

Fig. 34. — Diagram of tracheal system in body of beetle, sp., spiracles; tr., tracheae. 

(After Kolbe.) 
Fig. 35. — Diagram showing main tracheae in respiratory system of locust, Dissosteira 

Carolina. (After Snodgrass; twice natural size.) • 

Fig. 36. — Diagram showing respiratory system in thripf. St., spiracles. (After Uzel; 

much enlarged.) 

The Structure and Special Physiology of Insects 1 9 

on it not merely to take up oxgyen from the outer air and give up the 

waste carbon dioxide of the 
body, but also to convey these 
gases to and from all the tis- 
sues of the body. The blood 
is not red, but pale yellowish 
or greenish, and is really more 
like the lymph of the ver- 
tebrate body than like its 

Insects do not breathe 
through the mouth or any 
openings on the head, but have ' 
a varying number (usually 
from two to ten pairs) of 
small paired openings on the 
sides of the thorax and abdo- 
men. These openings, called 
spiracles, or stigmata, are ar- 
ranged segmentally and in 
most insects are to be found 
on two of the thoracic seg- 
ments and on all the abdomi- 
nal segments except the last two or three. The openings are euarded by fine 
hairs or even little valvular lids to prevent 
the ingress of dust, and are the entrances to 
an extended system of delicate air-tubes or 
tracheae which branch and subdivide until 
the whole of the internal body is reached 
and ramified by fine capillary vessels bring- 
ing fresh air to all the tissues and carrying 
off the waste carbon dioxide made by the 
metabolism of these tissues. The usual 
general arrangement of this elaborate re- 
spiratory system is shown in Figs. 34, 35, 
and 36. Short broad trunks lead from 
each spiracle to a main longitudinal trunk 
on each side of the body, from which _^^^m. 

numerous branches arise, these going to iS^^! 

particular regions of the body (Fig. 38) Fig. 39— i'lcce of trachea (air-tube), 
, , , , . » . ji .M greatly magnified, showing spiral 

and there branchmg \epeatedly until e^^^^j (t^nidia). (Photomicro- 

even individual cells get special tiny graph by George O. Mitchell.) 

Fig. 37 

Fig. 37. — Diagram showing respiratory system of pupa 
of mealy -winged fly, Akyrodes sp.; only two pairs 
of spiracles are present. (After Bemis; much 

Pig 28. — Diagram of trachese in head of cockroach. 
Note branches to all mouth-parts, and the an- 
tennae. /., tracheae, or air-tubes. (After Miall 
and Denny.) 

20 The Structure and Special Physiology of Insects 

respiratory capillaries. The tracheae are readily recognized under the micro- 
scope by their finely transversely ringed or striated appearance (Fig. 39). 
These transverse "rings" are really spirally arranged short chitinized 
thread-like thickenings on the inner w^all of the tube, which by their elasticity 
keep the delicate air-tubes open. The tubes are filled and emptied by a 

rhythmic alternately contracting and expanding 
movement of the abdomen, called the respiratory 
movement. When the ring-muscles contract, the 
walls of the abdomen are squeezed in against 
the viscera, which, compressing the soft air-tubes, 
force the air out of them through the spiracles; 
when the body-walls are allowed to spring back 
to normal position fresh air rushes in through the 
spiracles and fills up the air-tubes, which expand 
because of the elastic spiral thickenings in their 
walls. Insects which live in water either come 
up to the surface to breathe and in some cases 
to take down a supply of air held on the outside 
of the body by a fine pubescence like the pile of 
velvet, or they are provided with tracheal gills 
(Fig. 40) which enable them to breathe the air 
mixed with, or dissolved in, the water. Gillcd 
insects do not, of course, have to come to the 
surface to breathe. The gills may be thin plate- 
like flaps on the sides or posterior tip of the 
body, or may be tufts of short thread-like tubes 
variously arranged over the body. Or they 
may be, as in the dragon-fly nymphs, thin folds along the inner wall of the 
rectum, the water necessary to bathe them being taken in and ejected again 
through the anal opening. In all cases these insect gills differ from those 
of other animals, as crabs and fishes, in that they are not organs for the 
purification of the blood, i.e., effecting an exchange of carbon dioxide and 
oxygen carried by it, but are means for an osmotic exchange of the fresh 
air dissolved in water for carbon-dioxide-laden air from air-tubes or tracheae 
which run out into the gills. Probably no more blood enters these gills 
than is necessary to bring food to them. Impure air is brought to them 
by air-tubes, and exchanged by osmosis through the thin walls of air-tube 
and giU-membrane for fresh air, which passes from these gill air-tubes to 
the rest of the respiratory system of the body. 

The nervous system of insects shows the fundamentally segmental make-up 
of the body better than any of the other systems of internal organs, although 
probably in the successive chambers of the dorsal vessel or heart, and certainly 

Fig. 40. — Young (nymph) of 
May-fly showing {g.) tra- 
cheal gills. (After Jenkins 
and Kellogg.) 

The Structure and Special Physiology of Insects 21 

in the paired arrangement of the spiracles and tracheal trunks leading from 
them, a segmental condition is obvious. The central nervous system consists 




saLgl . "-,, 
i.b.wg.- •-// 
br. _ I' 


i.bjnsi. -E 

"''' i.b.h.-::.:::ff 



prov. iw^ 


susp. /fl- -- 

9'C- - |-|^ 


vent jfl 


ir. 11/ 

ad.tis yF\ 

ma I. tub. yi^:^--- 

FiG. 41. — Larva of giant crane-fly, Holorusia riibiginosa. A, entire; B, dissected, show- 
ing all organs except the muscles and ventral nerve-chain, h., head; ant., anterina; 
i.b.res., imaginal bud of pupal respiratory tube; i.b.wg., imaginal bud of wing;, imaginal bud of mesothoracic leg; i.b.h., imaginal bud of balancer;, imaginal bud of metathoracic leg (the imaginal buds of fore legs are con- 
cealed by head-capsule);, salivary gland (the other salivary gland is removed); 
br., brain; ces., oesophagus; prov., proventriculus; susp., suspensorium; g.c, gastric 
coecum; vent., ventriculus; tr., trachea; ad.tis., adipose tissue; mal.tub., Malpi- 
ghian tubule; d.v., dorsal vessel; w.m., wing-muscles of pericardium;, 
small intestine; tes., testis; ini.c, intestinal csecum; v.d., vas deferens; Lint., large 
intestine; sp., spiracle;, terminal processes. (Twice natural size.) 

of a brain and a ventral chain of pairs of ganglia segmentally arranged and 
connected by a pair of longitudinal cords or commissures (Figs. 42, 43, 44). 
The two members of each of the pairs of ganglia as well as of the pair of 

22 The Structure and Special Physiology of Insects 

Fig. 42. Fig. 43. Fig. 44. 

Fig. 42. — Diagram of ventral nerve-cord of locust, Dissosteira Carolina. (After Snod- 

grass; twice natural size.) 
Fig. 43. — Diagram of the nervous system of the house-fly. (After Brandt; much 

Fig. 44. — Nervous system of a midge, CJuronomus sp. (After Brandt, much enlarged.) 

commissures are in most insects more or less fused to form single ganglia 
and a single commissure, but in others the commissures, 
at least, are quite distinct. In the simpler or more 
generalized condition of the nervous system as seen 
in the simpler insects and the larvae of the higher 
ones there are from three or four to seven or eight 
abdominal ganglion pairs, one pair to a segment, a 
pair in each of the three thoracic segments, and one 
in the head just under the oesophagus. From this 
ganglion (or fused pair) circumoesophageal commis- 
sures run up around the oesophagus to an important 
ganglion (also composed of the fused members of a 
pair) lying just above the oesophagus and called the 
brain, or supraoesophageal ganglion (Figs. 45, 46, and 
47). From this proceed the nerves to those impor- 
tant organs of special sense situated on the head, the 
antennae and eyes. From the suboesophageal gan- 
lion nerves run to the mouth-parts, from the thoracic ganglia to the 

Fig. 45. — Brain, com- 
pound eyes, and part 
of sympathetic nerv- 
ous system of locust, 
Dissosteira Carolina. 
(After Snodgrass; 
greatly magnified.) 

The Structure and Special Physiology of Insects 23 

wings and legs and the complex thoracic muscular system, while from 
the abdominal ganglia are innervated the abdominal muscles and sting, 
ovipositor, or male claspers. In addition to this main or ventral nervous 
system there is a small and considerably varying sympathetic system (Figs. 
46 and 48) to which belong a few minute ganglia sending nerves to those 
viscera which act automatically or by reflexes, as the alimentary canal and 
heart. This sympathetic system is connected with the central or principal 

Fig. 46. 

Fig. 46. — Brain, circumcesophageal commissures, and subcesophageal ganglion of the 
red-legged locust, Melanophis jemur-rubrum. oc, ocellus; op.n., optic nerve; a.n., 
antennal nerve; m.oc, middle ocellus; op.I., optic lobe; a.L, olfactory lobe; a.s.g., 
anterior sympathetic ganglion; p.s.g., posterior sympathetic ganglion; f.g., frontal 
sympathetic ganglion; Ibr., nerve to labrum; oe.c, circumoesophageal commissure; 
g*, subcesophageal ganglion; md., nerve to mandible; mx., nerve to maxilla; l.n., 
nerve to labium; «., unknown nerve, perhaps salivary. (After Burgess; greatly 

Fig. 47. — Cross-section of brain, oesophagus, circumcesophageal commissures, and 
subcesophageal ganglion of larva of the giant crane-fly, Holoriisia rubiginosa. 

nervous system by commissures which meet the brain just at the origin 
from it of the circumoesophageal commissures. 

The specialization of the ventral nerve-chain is always of the nature of 
a concentration, and especially cephalization of its ganglia (Figs. 49 and 
50). The abdominal ganglia may be fused into two or three or even into 
one compound ganglion; or indeed all of them may migrate forward and 
fuse with the hindmost thoracic ganglion, thus leaving the whole abdomen 

24 The Structure and Special Physiology of Insects 

to be innervated by long nerves running from the thorax. The thoracic 
gangha may fuse to form one, and in extreme cases all the abdominal and 
thoracic ganglia may be fused into one large mid- 
thoracic center. 

In tracing the development of the nervous 
system during the ontogeny of one of the special- 
ized insects, the changes from generalized condi- 
tion, i.e., presence of numerous distinct ganglia 
segmentally disposed, shown in the newly hatched 



48. riG 49. 

Fig. 48. — Part of sympathetic nervous system of larva of harlequin-fly, Chironomus 
dorsalis. oes., oesophagus; j.g., frontal ganglion; r.n., recurrent nerve; d.v., dorsal 
vessel; n*, nerve passing from brain to frontal ganglion (Newport's fourth nerv-e); 
hr., brain; rn., point of division of recurrent nerve; tr., tracheae; pg., paired ganglia; 
d.v.Ji., nerve of dorsal vessel; d.v.g., ganglia of dorsal vessel; g.n., gastric nerve 
to cardiac chamber. The course of the recurrent nerve beneath the dorsal vessel is 
dotted. (After Miall and Hammond; greatly magnified.) 

Fig. 49. — Stages in the development of the nervous system of the honey-hee, Apis nielli- 
fica; I showing the ventral nerve-cord in the youngest larval stage, and 7 the system 
in the adult. (After Brandt; much enlarged.) 

larva, to specialized condition, i.e., extreme concentration and cephalization, 
that is, migration forward and fusion of the ganglia, shown in the adult, 
are readily followed (Figs. 49 and 50). 

The special senses of insects and the sense-organs are of particular inter 
est because of the marked unusualness of the character of the specialization 
of both the organs and senses, as compared with the more familiar condi- 
tions of the corresponding organs and functions of our body. The world 
is known to animals only by the impressions made by it on the sense-organs, 

The Structure and Special Physiology of Insects 25 

and the particular condit'on of functioning of these organs, therefore, is of 
unique importance in the life of any particular animal. If the senses vary 
much in their capacities among different animals, the world will have a differ- 
ent seeming to different creatures. It will be chiefly known to any par- 
ticular species through the dominant sense of that species. To the con- 
genitally blind the world is an experience of touched things, of heard things, 
and of smelled and tasted things. To the bloodhound it is known chiefly 
by the scent of things. It is a world of odors; the scent of anything deter- 
mines its dangerousness, its desirableness, its interestingness. As insects 
know it, then, the world depends largely upon the particular character and 
capacity of iheir sense-organs, and we reahze on even the most superficial 
examination of the structure of these organs, and casual observation of the 

Fig. 50. — Stages in the development of the nervous system of the water-beetle, ^cilins 
sulcatus; i showing the ventral nerve-cord in the earliest larval stage, and 7 the 
system in the adult. (After Brandt; much enlarged.) 

responses of insects to those stimuli, like sound-waves, light-waves, dis- 
solved and vaporized substances, which affect the sense-organs, that the 
insects have some remarkable special sense-conditions. But the difficul- 
ties in the way of understanding the psychology of any of the lower animals 
are obvious when it is recalled that our only knowledge of the character 
of sense-perceptions has to depend solely on our experience of our own per- 
ceptions, and on the basis of comparison with this. We do not know if 
hearing is the same phenomenon or experience with insects as with us. 
But a comparison of the morphology of the insect sense-organs with that 
of ours, and a course of experimentation with the sight, hearing, smelling, 
etc., of insects, based on similar experimentation with our own senses, leads 
us to what we believe is some real knowledge of the special sense-condi- 
tions of insects. 

2 6 The Structure and Special Physiology of Insects 

Insects certainly have the senses of touch, hearing, taste, smell, and sight. 
If they have others, v^e do not knovv^ it, and probably cannot, as we have 

no criteria for recognizing others. 
The tactile sense resides especially 
in so-called "tactile hairs," scattered 
more or less abundantly or regu- 
larly over the body. Each of these 
hairs has at its base a ganglionic 
nerve-cell from which a fine nerve 
runs to some body ganglion (Fig. 51). 
They are specially numerous and 
conspicuous on the antennas or 
" feelers," and often on certain pro- 
cesses called cerci, projecting from 
the tip of the abdomen. They may 
occur, however, on any part of the 
body, and are usually recognizable 
by their length and semi-spinous nature. The sense of taste resides 
in certain small papillae, usually two-segmented, or in certain pits, which 

Fig. 51. — Diagram showing innervation of a 
tactile hair, sh., tactile hair; ch., chitinized 
cuticle; hyp., hypoderm, or cellular layer 
of the skin; s.c, ganglion cell; c.o., gan- 
glion of the central nervous system. (After 
vom Rath.) 

Fig. 52. Fig. 53. 

Fig. 52. — Nerve-endings in tip of maxillary palpus of Locusta viridissima. s.h., sense- 
hairs; i'.c, sense-cells; 6.c., blood-cells. (After vom Rath; greatly magnified.) 

Fig. 53. — -Nerve-endings in tip of labial palpus of Machilis polypoda. (After vom 
Rath; greatly magnified.) 

occur on the upper wall of the mouth (epipharynx) and on the mouth- 
parts, especially the tips of the maxillary and labial palpi, or mouth- 
feelers. As substances to be tasted have to be dissolved, and have to 

The Structure and Special Physiology of Insects 27 

come into actual contact with the special taste nerves, it is obvious 
that insects, to taste solid foods, have first to dissolve particles of these 
foods in the mouth-fluids, and that the taste-organs have to be situated 
in the mouth or so that they can be brought into it to explore the food, as 
are the movable, feeler-like palpi. What experimentation on the sense of 
taste in insects has been carried on shows that certain insects certainly taste 
food substances, and indicates that the sense is a common attribute of all 
insects. Lubbock's many experiments with ants, bees, and wasps present 
convincing proof of the exercise of the taste sense by these insects. Forel 
mixed morphine and strychnine with honey, which ants, attracted by the 
honey smell, tasted and refused. Will's experiments show that wasps 
recognize alum and quinine by taste. He found bees and wasps to have 
a more delicate gustatory sense than flies. 

Smell is probably the dominant special sense among insects. It exists 
at least in a degree of refinement among certain forms that is hardly 
equalled elsewhere in the animal kingdom. The smelling organs are micro- 
scopic pits and minute papillte seated usually and especially abundantly 
on the antennae, but probably also occurring to 
some extent on certain of the mouth-parts. The 
fact that the antennae are the principal, and in 
many insects the exclusive, seat of the olfactory 
organs has been proved by many experiments in 
removing the antennae or coating them with par- 
affine. Insects thus treated do not find food or 
each other. As substances to be smelled must, 
actually come into contact, in finely divided con- 
dition, with the olfactory nerve-element, these 
pits and papillae are arranged so as to expose 
the nerve-end and yet protect it from the 
ruder contact with obstacles against which the 
antennae may strike. It is certain that most 
insects find their food by the sense of smell, and 
the antenna of a carrion-beetle (Fig. 54) shows 
plainly the special adaptation to make this sense 
highly effective. On the "leaves" of each antenna 
of June-beetles nearly 40,000 olfactory pits occur. 
Some of the results of experimentation on smell 
indicate a delicacy and specialization of this sense 
hardly conceivable. A few examples will illustrate 
this. It is believed that ants find their way back 
to their nests by the sense of smell, and that 
they can recognize by scent among hundreds of individuals taken from 

Fig. 54. —Antenna of a 
carrion-bcctle with the 
terminal three segments 
enlarged and flattened, 
and bearing many smell- 
ing-pits. (Photomicro- 
graph by George O. Mit- 
chell; much enlarged.) 

2 8 The Structure and Special Physiology of Insects 

various communities the members of their own community. Miss Fielde's 
experiments show that the recognition of ants by each other depends on the 
existence of a sense of smell of remarkable differentiative capacity. The 
odors of the nest, of the species, of the female parent, and of the individ- 
ual are all distinct and perceivable by the smelling-organs, situated on 
distinct particular antennal segments. In the insectary at Cornell University 
a few years ago a few females of the beautiful large promethea moth were 
put into a covered box which was kept inside of the insectary building. 
No males of this moth species had been seen about the insectary nor in 

its immediate vicin- 
ity for several days, 
although they had 
been specially sought 
for by collectors. 
Yet in a few hours 
after the female 
moths were first con- 
fined nearly fifty 
male prometheas 
were fluttering about 
outside over the glass 
roof of the insectary. 
They could not see 
the females, but un- 
doubtedly discovered 
them by the sense of 
smell. These pro- 
methea moths have 
elaborately branched 
or feathered anten- 
nae, affording area 
for very many smell- 

Mayer's experiments with promethea also reveal the high specialization 
of the sense of smell. This investigator carried 450 promethea cocoons 
from Massachusetts to the Florida keys. Here on separated small 
islands the moths issued from the cocoons, hundreds of miles south of their 
natural habitat. This isolation insured that no other individuals than 
those controlled by the experimenter could confuse the observations. 
Female moths w^re confined in glass jars with the mouth closed by 
netting. Other females were confined in smaller glass jars turned upside 
down and the mouth buried in sand. Males being released at various 

Fig. 55. — Auditory organ of a locust, Melanoplus sp. The 
large clear part in the center of the figure is the thin tym- 
panum with the auditory vesicle (small, black, pear-shaped 
spot) and auditory ganglion (at left of vesicle and connected 
with it by a nerve) on its inner surface. (Photomicrograph 
by George O. Mitchell; greatly magnified.) 

The Structure and Special Physiology of Insects 29 

distances soon found their way to the jar (containing females) which had 
its mouth open to the air, but no male came to the jar with its mouth her- 
metically sealed. Through the glass sides of both 
jars the females were plainly visible. The antenna; 
of certain males were covered with shellac. These 
males, when released, never found the females, and 
often paid no attention to them when brought within 
an inch of their bodies. Of other males the eyes 
were covered with pitch; but these males had no 
difficulty whatever in finding the females. It is 
plainly obvious from these experiments that the 
males found the females wholly by scent and not at 
all by sight. 

That some insects hear is proved by their posses- 
sion of auditory organs, and has also been demon- Fig. 56.— Male mos(|uito, 
strated bv experiment. The fact, too, that manv f o^i"g '^«-^'-) antennal 

,'. , . ■' hairs. (After Jordan 

msects have special sound-makmg apparatus and and Kellogg; three times 
do make characteristic sounds is a kind of proof natural size.) 
that they can also hear. The auditory organs of insects, curiously enough, 
are of several kinds and are situated on different parts of the body, in 

various species. Among the locusts, 
katydids, and crickets, the most con- 
spicuous of all the sound-making in- 
sects except the cicada, the ears are 
small tympanic membranes on the 
base of the abdomen in the locusts 
(Fig. 55), and on the tibiae of the fore 
legs in the katydids and crickets. 
Associated with each tympanum is a 
small liquid-filled vesicle and a special 
auditory ganglion from which an 
auditory nerve runs to one of the 
ganglia of the thorax. Among the 
Fig. 57.— Diagram of longitudinal section midges and mosquitoes the antennae- 
through first and second antennal seg- those all-important sensitive Structures 

ments of a mosciuito, Mochlonyx ciilici- , , ,, • i i • i 

formis, male, showing complex auditory ~^^^ abundantly provided With cer- 

organ composed of fine chitinous rods, tain fine long hairs, the auditory hairs 
nerve-fibers, and nerve-cells. (After /tt- _/:\ u* u ^ i tu ' j 

Child; greatly magnified.) ^^^S- S6), which take up the SOUnd- 

waves and transmit the vibrations to an 
elaborate percipient structure composed of many fine chitin-rods and ganglion- 
ated nerves contained in the next to basal antennal segment (Fig. 57). From 
this segment runs a principal auditory nerve to the brain. Many other insects 

30 The Structure and Special Physiology of Insects 

besides the midges and mosquitoes possess this type of auditory organ; 
in fact such an organ, more or less well developed, has been found in almost 
every order except the Orthoptera (the order of locusts, crickets, katydids, 
etc.) in which the tympanic auditory organs occur. 
Special isolated hairs scattered sparsely over the 
body, connected with a special peripheral nervous 
arrangement, are believed by some entomologists 
to be a third kind of auditory structure, and are 
called chordotonal organs. Experimentally the 
sense of hearing has been surely determined for 
certain insects. A single striking example of this 
experimentation must here suffice. Mayer fastened 
a live male mosquito to a glass slide, put it under 
a microscope, and had a series of tuning-forks of 
different pitch sounded. When the Ut^ fork of 
512 vibrations per second was sounded many of 
the antennal hairs were set, sympathetically, into 
strong vibration. Tuning-forks of pitch an octave 
lower and an octave higher also caused more 
vibration than any intermediate notes. The male 
mosquito's auditory hairs, then, are specially fitted to respond to, i.e., be 
stimulated by, notes of a pitch produced by 512 vibrations. Other, but 
fewer, hairs of different length vibrated in response to other tones. Those 
auditory hairs are most affected which are at right angles to the direction 
from which the sound comes. From this it is obvious that, from the position 
of the antennae and the hairs, a sound will be loudest or most intense if it is 
directly in front of the head. If the mosquito is attracted by sound, it will 
thus be brought straight head end on toward the source of the sound. As a 

Fig. 58. — Longitudinal sec- 
tion through ocellus of the 
honey-bee, Apis mellifica. 
/., cuticular lens; i.e., cell- 
ular layer of skin; c.b., 
crystalline layer; r.c, ret- 
inal cells; o.n., optic 
nerve. (After Redikor- 
zew; greatly magnified.) 

Fig. 50. — OccUar lens of larva of a saw-fly, Cimhex sp., showing its continuity with the 
chitinized cuticle. (After Redikorzew; greatly magnified.) 

matter of fact, Mayer found the female mosquito's song to correspond nearly 
to Ut4, and that her song set the male's auditory hairs into vibration. With 
little doubt, the male mosquitoes find the females by their sense of hearing. 

Insects have two kinds of eyes, simple and compound. On most 
species both kinds are found, on some either kind alone, and in a few no 
eyes at all. Blind insects have lost the eyes by degeneration. The most 

The Structure and Special Physiology of Insects 3 i 

Fig. 60. — Part of corneal cuti- 


primitive living insects, Campodea and otliers, have eyes, although only 

simple ones. The larva; of the specialized 
insects, i.e., those with complete metamor- 
phosis, also have only simple eyes. The com- 
pound eyes are not complex or specialized 
derivations of the simple ones, but are of in- 
dependent origin and of obviously distinct 
structural character. The simple eyes, also 
called ocelli (Fig. 58), which usually occur to 
the number of three in a little triangle on 
top of the head, are small and inconspicuous, 
and consist each of a lens, this being simply 
cle, showing facets, of the ^ small convexly thickened clear part of the 
fl^?^^rt/.S5fp.^Photo- chitinized cuticle of the head-wall (Fig. 59) 
micrograph by George O. and a group of modified skin-cells behind it 
Mitchell; greatly magnified.) gp^.j^^y provided with absorbent pigment and 

capable of acting as a simple light-sensitive or retinal 
surface. The ocellus is supplied with a special nerve 
from the brain. The compound eyes are always 
paired and situated usually on the dorso-lateral parts 
of the head; they are usually large and conspicu- 
ous, sometimes, as in the dragon-flies and horse- 
flies, even forming two-thirds or more of the mass 
of the head. Externally each compound eye pre- 
sents a number (which varies all the way from a 
score to thirty thousand) of facets or microscopic 
polygonal cuticular windows (Fig. 60). These are 
the cornea of the eye. Behind each facet is a dis- 
tinct and independent subcylindrical eye-element or 
ommatidium composed of a crystalline cone (want- 
ing in many insects) enveloping pigment (which pre- 
sumably excludes all light-rays except those which 
fall perpendicularly or nearly so to the corneal 
lens of that particular ommatidium), and a slender y(|lir~--OIl 

tapering part including or composed of the nervous Fig. 61.— Longitudinal 

or retinal element called rhabdom (Fig. 61). Each f^tion through a few 

. • 1 1 • r 1 facets and eye-elements 

of these ommatidia perceives that bit of the external (ommatidia) of the 

object which is directly in front of it; i.e., from which compound eye of a 

light is reflected perpendicularly to its corneal facet. "j°. ' /^' ^°crysulHne 

All of these microscopic images, each of a small part cones; p., pigment; r., 

of the external object, form a mosaic of the whole ^erve * ^"(ffter' ExSJ-'j 

object, and thus give the familiar name mosaic greatly magnified.) 

32 The Structure and Special Physiology of Insects 

vision to the particular kind of seeing accomplished by the compound 

The character or degree of excellence of sight by the two kinds of 
eyes obviously varies much. The fixed focus of the ocelli is extremely short, 


FiG. 62. 

Fig. 64. 


Fig. 62. — Longitudinal sections through outer part of eye-elements (ommatidia) of com- 
pound eyes of Lasiocampia quercijolia; ommatidia at left showing disposition of 
pigment in eyes in the light, at right, in the dark. (After Exner; greatly magnified.) 

Fig. 63. — Longitudinal section through a few eye-elements of the compound eye of Cato- 
cola niipta; left ommatidia taken from an insect killed in the dark, right ommatidium 
taken from insect killed in the light. (After Exner; greatly magnified.) 

Fig. 64. — Section through the compound eyes of a male May-fly, showing division of 
each compound eye into two parts, an upper part containing large eye -elements 
(ommatidia), and a lower part containing small eye-elements (ommatidia). (.\fter 
Zimmerman; greatly magnified.) 

and probably the range of vision of these eyes is restricted to an inch or 
two in front of the insect's head. Indeed entomologists commonly believe 
that the ocelli avail little beyond distinguishing between hght and darkness. 
With the compound eyes the focus is also fixed, but is longer and the range 
of vision must extend to two or three yards. It is obvious that the larger 

The Structure and Special Physiology of Insects 33 

and more convex the eyes the wider will be the extent of the visual field, 
while the smaller and more abundant the facets the sharper and more dis- 
tinct will be the image. Although no change in focus can be effected, cer- 
tain accommodation or flexibility of the seeing function is obtained by the 
movements of the pigment (Figs. 62 and 63) tending to regulate the amount 
of light admitted into the eye (as shown by Exner), and by a difference in size 
and pigmental character of the ommatidia (Fig. 64) composing the com- 
pound eyes of certain insects tending to make part of the eye especially 

Fig. 65. — A section through the compound eye, in late pupal stage, of a blow-fly, Calli- 
phora sarracenicp. In the center is the brain with optic lobe, and on the right-hand 
margin are the many eye-elements (ommatidia) in longitudinal section. (Photomi- 
crograph by George O. Mitchell; greatly magnified.) 


adapted for seeing objects in motion or in poor light, and another part for 
seeing in bright light and for making a sharper image (as shown by Zim- 
merman for male May-flies, and by myself for certain true flies (see p. 318)). 
Our careful studies of the structure of the insect eye, and the experimentation 
which we have been able to carry on, indicate that, at best, the sight of 
insects cannot be exact or of much range. 

The psychology of insects, that is, their activities and behavior as deter- 
mined by their reflexes, instincts, and intelligence, is a subject of great inter- 
est and attractiveness, but obviously one difficult to study exactly. The 

34 The Structure and Special Physiology of Insects 

elaborateness of many insect instincts, such as those of the ants, wasps, and 
bees, to choose examples at once familiar and extreme in their complexity, 
makes it very difficult to analyze the trains of reactions into individual ones, 
and to determine, if it is indeed at all determinable, the particular stimuli 
which act as the springs for these various reactions. The attitude of the 
modern biologist in this matter would be to keep first in mind the theory 
of reflexes, to look keenly for physico-chemical explanations of the reac- 
tions, and only when forced from this position by the impossibility of find- 
ing mechanical explanations for the phenomena to recognize those com- 
plex reflexes which we call instincts, and finally those acts which we call 
intelligent, or reasonable, and which are possible only to the possessors of 
associative memory. The investigations, mostly recent, which have been 
directed toward a determination of the immediate springs or stimuli of 
insect reactions indicate clearly that many of these responses, even some 
which were formerly looked on as surely indicative of considerable intelli- 
gence on the part of their performers, are explicable as rigid reflex (mechan- 
ical) reactions to light, gravity, the proximity of substances of certain 
chemical composition, contact with solid bodies, etc. On the other hand 
the position of the extreme Upholders (Bethe, UexkuU, and others) of the 
purely reflex explanation of all insect behavior will certainly prove untenable. 
As one of the phases of insect biology to which this book is particularly 
devoted is that which includes the study of habits, activities, or behavior, 
we may dispense with any special discussion of instinct in this introductory 
chapter. It is sufficient to say that no other class of invertebrate animals 
presents such an interesting and instructive psychology as the insects. 




HAT animals are born or hatch from eggs in 
an immature condition is such famihar natural 
history that we are likely to overlook the 
significance and consequences of the fact unless 
our attention is particularly called to them. 
This condition of immaturity makes it necessary 
that part of the free life of the organism has 
to be devoted to growth and development and 
has to be undergone in an imperfect condition, 
a condition of structure and physiology, indeed, which may be very different 
from that of the parents or of maturity. While most animals that are born 
alive re:emble the parents in most respects, always excepting that of size, 
many of those animals which hatch from eggs deposited outside the body 
of the mother issue from the egg with few indeed of the characteristics of the 
parents and may be so dissimilar from them that only our knowledge of 
the life-history of the animal enables us to recognize these young individuals 
as of the same species as the parent. The butterfly hatching as the worm- 
like caterpillar, and the frog as the fish-like tadpole, are the classic examples 
of this phenomenon. The mammals, our most familiar examples of animals 
which give birth to their young alive and free, nourish, for weeks or months 
before birth, the developing growing young. But with egg-laying animals 
usually only such nourishment is furnished the young as can be enclosed 
as food-yolk within the egg-shell. As a matter of fact, some young which 
hatch from eggs, as, for example, chickens, quail, etc., hatch in well- 
developed condition; and some young mammals, nourished by the mother's 
body until birth, are in a conspicuously undeveloped state, as a young 
kangaroo or opossum. But nevertheless it is generally true that an animal 
hatched from an egg has still a larger amount of development to undergo 
before it comes to the stature and capacity of its parents than one which is 



Development and Metamorphosis 

born alive, after having passed a considerable time growing and developing 
in the body of the mother. And this difference in degree of development at 
birth is largely due simply to the difference in amount of nourishment 
which can be afforded the young. The embryo in the egg uses up its food 
early in its developmental career and before it has reached the stage of 
likeness to its parents. It issues in a condition picturing some far-distant 
ancestor of its species, or more frequently, perhaps, in a modified, adapted 
condition, fit to make of this tender unready creature thus thrust before 
its time into the struggle for living an organism capable of caring for itself, 
although not yet endowed with capacities as effective as, or even similar to, 
those of the parent. 

It is familiar to us, then, that development is not wholly postnatal or 
postembryonic ; that before birth or hatching a greater or less amount of 

development, requiring a longer or shorter 
period of time, has already been undergone. 
Every animal begins life as a simple cell; all 
animals except the Protozoa (the simplest ani- 
mals, those whose whole body for its whole 
life is but a single cell) finish life, if red 
Nature permits them to come through myriad 
dangers safely to maturity, as a complex of 
thousands or millions of cells united into 
great variety of tissues and organs. This 
great change from most simple to most complex 
condition constitutes development: the actual 
increase of body-matter and extension of 
dimensions is growth. 

Most insects hatch from eggs; being born 
alive is the exceptional experience of the young 
of but few kinds, and even this is a sort of 
pseudo-birth. Such hatch alive, one may better 
say, for they begin life in eggs, not laid out- 
side the mother body to be sure, but held in 
the egg-duct until hatching-time. With very few exceptions, young insects 
are not nourished by the mother except in so far as she stores a supply of 
yolk around or by the side of each embryo inside the egg-shell. The form- 
ing of the egg is a matter which does not lend itself readily to the observa- 
tion and study of amateurs, but is a phenomenon of unusual interest to 
whomever is privileged to discover it. The insect ovaries consist of a pair 
of little compact groups of short tapering tubes (Fig. 66). In the anterior or 
beginning end of each tube is a microscopic space or chamber from whose 
walls cells loosen themselves and escape into the cavity. These cells become 

Fig. 66. — Ovaries and oviducts 
of a thrips. o./., ovarial tubes; 
o.d., oviduct; r.s., seminal 
receptacle, or spermatheca; 
d.r.s., duct of the seminal re- 
ceptacle. (After Uzel; much 

Development and Metamorphosis 


either the germinal or the food part of the eggs. There seems to exist no 
dififerentiation among these cells at first, but soon certain ones begin to 
move slowly down through the egg-tube in single file, each becoming sur- 
rounded and enclosed by yolk, i.e., reserve foodstufi^. This gathering of 
yolk increases the size of the forming eggs, so that they appear as a short 
string of beads of varying size enclosed in the elastic egg-tube. When of 
considerable size each egg in the lower end of the tube becomes enclosed 



Fig. 67. — Insect eggs and parts of eggs, showing micropyle. a, egg of Drosophila cel- 
laris; h, upper pole of egg of robber-fly, Asilus crabriformis; c, upper pole of egg 
of hawk-moth, Sphinx popiili; d, egg of head-louse, Pediculus capitis; e, egg of 
dragon-fly, Libellida depressa; /, upper surface of egg of harpy-moth, Harpyia 
vinida; g, upper pole of egg of Hammalicherus cerdo; h, upper pole of egg of sul- 
phur-butterfly, Colias hyale. (After Leuckart; much enlarged.) 

in two envelopes, a membranous inner one (yolk or vitelline membrane) and 
an outer horny one, the chorion or egg-shell. But both of these envelopes 
are pierced at one pole by a tiny opening, the micropyle (Fig. 67), and 
through this opening the fertilizing spermatozoa enter the egg from the 
seminal receptacle just before the egg is extruded from the body. 

The development of the embryo within the egg is also securely sealed 
away from the eyes of most amateurs. The study of insect embryology 
requires a knowledge of microscopic technic, and facilities for fixing and 


Development and Metamorphosis 

imbedding and section-cutting which are not often found outside the college 
laboratory. But the particularly interesting and suggestive stages in this 
development may be outlined and illustrated in brief space. First, the 
germinal cell near the center of the egg divides repeatedly (Fig. 68 A) and 
the resulting new cells migrate outward against the inner envelope of the 
egg and arrange themselves here in a single peripheral layer, called the 
blastoderm (Fig. 68 D, bl). On what is going to be the ventral side of the 
egg the cells of the blastoderm begin to divide and mass themselves to form 
the ventral plate (Fig. 69 C). The embryo is forming here; the rest of the 
blastoderm becomes modified and folded to serve as a double membranous 
envelope (called amnion and serosa) for the embryo. Stretching nearly from 
pole to pole as a narrow streak along the ventral aspect of the egg, the 


Fig. 68. — Early stage in development of egg of water-scavenger beetle, Hydrophilus sp. 
A, first division of nucleus; B, migration of cleavage-cells outward; C, beginning 
of blastoderm; D, blastoderm; y., yolk; dc, cleavage-cells; yc, yolk-cells; bl., 
blastoderm. (After Heider; greatly magnified.) 

developing embryo begins soon to show that fundamental structural charac- 
teristic of insects, a segmental condition (Fig. 6gD). One can now make 
out the forming body-rings or segments, and each soon shows the beginnings 
or rudiments of a pair of appendages (Fig. 6gE). The appendages of the 
head and thoracic segments continue to develop and begin soon to assume 
their definitive character of antennae, mouth-parts, and legs, but those of the 
abdominal segments never get farther than a first appearance and indeed 
soon disappear. In the mean time the internal systems of organs are grad- 
ually developing, the ventral nerve-chain first, then the alimentary canal, 
and later the muscles, tracheae, and the heart. All the time the yolk is 
being gradually used up, fed on, by the cells of the developing and growing 
embryo, until finally comes the disappearance of all the stored food, and the 
time for hatching. 

Development and Metamorphosis 


The eggs have been laid, because of the remarkable instinct of the 
mother, in a situation determined chiefly by the interests of the young 
which are to hatch from them. The young of many kinds of insects take 
very different food from that of the mother — a caterpillar feeds on green 
leaves, the butterfly on flower-nectar — or live under very different circum- 
stances — young dragon-flies and May-flies live under water, the adults in 
the air. A monarch butterfly, which does not feed on leaves, nor has ever 
before produced young, seeks out a milkweed to lay its eggs upon. The 
young monarchs, tiny black-and-white-banded caterpillars, feed on the 


Fig. 69. — Early stages in the development of the egg of saw-fly, Hylotoma beriheridis. 
C, ventral plate removed from egg; D, ventral plate, showing segmentation of body; 
E, embryo, showing developing appendages; F, same stage, lateral aspect; G, older 
stage, lateral aspect, ant., antenna; nid., mandible; mx., maxilla; li., labium; /', P, l^, 
legs; sg., salivary glands; St., spiracles; ab.ap., abdominal appendages; n.c, nerve- 
centers; a., anal opening; lb., labrum; sd., oesophageal invagination; y., yolk; 
b.s., abdominal segments; pd., intestinal invagination; am., amnion; s., serosa. 
(After Graber; greatly magnified.) 

green milkweed leaf-tissue; indeed they starve to death if they cannot have 
leaves of precisely this kind of plant! The reason that the butterfly, whose 
only food is the nectar of almost any kind of flower, ranges wide to find a 
milkweed for its eggs, is one not founded on experience or teaching or lea- 
son, but on an inherited instinct, which is as truly and as importantly an 
attribute of this particular species of butterfly as its characteristic color 
pattern or body structure. And the female of the great flashing strong- 
winged dragon-fly, queen insect of the air, when egg-laying time comes, 
feels a strange irresistible demand to get these eggs into water, dropping 
them in from its airy height, or swooping down to touch the tip of the abdo- 


Development and Metamorphosis 

men to the water's surface, there releasing them, or even crawling down 
some water-plant beneath the surface and with arduous labor thrusting the 
eggs into the heart of this submerged plant-stem. From the eggs hatch 
wingless dwarf-dragons of the pond bottom, with terrible extensile, clutch- 
ing mouth-parts and an insatiable hunger for living prey. 

So our young insects, after completing their embryonic development, 
come to the time of their appearance as free individuals compelled to find 
their own food and no longer sheltered by a firm egg-shell from the strenu- 

FiG. 70. — Series of stages in development of egg of fish-moth, Lepisma sp. A, begin- 
ning embryo; B, embryo showing segmentation; C, embryo showing appendages; 
D, embryo more advanced; E, embryo still more advanced; F, embryo still older 
and removed from egg; G, embryo removed from egg at time of readiness to hatch. 
y., yolk; emb., embryo; ser., serosa; am., amnion; ant., antenna; lb., labrum; 
md., mandible; mx., maxilla; mx.p., maxillary palpus; //., labium; H.p., labial 
palpus; /'.. P, P, legs; pr., proctodaeum, or intestinal invagination; cer., cerci; mp., 
middle posterior process. (After Heymons; greatly magnified.) 

ous fighting and hiding of the open road. Now these young insects, depend- 
ing upon how far they have carried their developmental course in the egg, 
hatch either almost wholly like their parents (excepting always in size), or 
in a condition fairly resembling the parents, but lacking all traces of wings 
and showing other less conspicuous dissimilarities, or finally they may appear 
in guise wholly unlike that of their parents, in such a condition indeed that 
they would not be recognized as insects of the same kind as the parents. 
But in all cases the young are certain, if they live their allotted days or weeks 

Development and Metamorphosis 


or months, to attain finally the parent structure and appearance. This 
attainment is a matter of further development, of postembryonic develop- 
ment, and the amount or degree of this development or change is obviously 
determined by the remoteness or nearness of the young at the time of hatch- 
ing to the adult or parental condition. The young of many of our most 
familiar insects, as beetles, flies, moths and butterflies, and ants, bees, and 
wasps, hatch out extremely unlike their parents in appearance: the well- 
known worm-like caterpillars of butterflies and moths are striking examples 
of this unlikeness. The changes necessarily undergone in the develop- 
ment from caterpillar to butterfly are so great that there actually results 
a very considerable degree of making over, or metamorphosis of the insect, 
and for convenience of roughly classifying insects according to their develop- 
ment, entomologists have adopted the terms complete metamorphosis, 
incomplete metamorphosis, and no metamorphosis to indicate three not 
very sharply distinguished kinds or degrees of postembryonic development. 

In the latter category are comparatively few species, because most insects 
have wings, and no insect is winged at birth. But the members of the sim- 
plest order (Aptera) are all primitively wingless, and their 
young are, in practically all particulars except body size and 
the maturity of the reproductive glands, like the adults 
(Fig. 71) ; their development may fairly be said to take place 
without metamorphosis. In addition to these primitively 
simple insects there are certain degenerate wingless species 
like the biting bird-lice, for example, whose young also 
reach the parental stature and character without meta- 

In the next category, that of development with in- 
complete metamorphosis, are included two large orders 
of insects and several smaller ones. All the sucking-bugs Y\ 
(order Hemiptera) and all the locusts, katydids, crickets, 
and cockroaches (composing the order Orthoptera), as well 
as the May-flies, dragon-flies, white ants, and several other 
small groups of unfamiliar forms, agree in having their 
young hatched in a condition strongly resembling the 
parents, although lacking wings, and in some cases, particu- 
larly those in which the young live on different food and in a different habitat 
from the adults, differing rather markedly in several superficial characters. 
Such is the case, for example, with the dragon-flies, whose young are aquatic 
and breathe by means of tracheal gills, and are provided with specially con- 
structed seizing and biting mouth-parts. But in such essential character- 
istics as number of legs, character of eyes and antennas, and, usually, char- 
acter of mouth-parts, the young and parent agree. During postembryonic 

I . — Young 
and adult of Po- 
dura sp., one of 
the simplest in- 
sects, showing 
without meta- 
(Much enlarged.) 


Development and Metamorphosis 

Fig. 72. — Developing stages, after hatching, of a locust, Melanoplus femur-ruhrumt 
a, just hatched, without wing-pads; h, after first moulting; c, after second moulting, 
showing beginning wing-pads; d, after third moulting; e, after fourth moulting, 
/, adult with fully developed wings. (After Emerton; younger stages enlarged; 
adult stage, natural size.) 

Fig. 73. — Stages in development of the wings of a locust. /., developing rudiment of 
fore wing; h., developing rudiment of hind wing; w., wing-pad. (After Graber; 
twice natural size.) 

Development and Metamorphosis 


development the young have to develop wings and make what other change 
is necessary to reach the adult type, but the life is continually free and active 
and the change is only a simple gradual transformation of the various parts 
in which differences exist. A common locust is an excellent example of 
an insect with such incomplete metamorphosis. Fig. 72 shows the develop- 
ing locust at different successive ages, or stages, as these periods are called 
because of their separation from each other by the phenomenon, common 
to all insects, of moulting. As the insect grows it finds its increase of girth 
and length restrained by the firm 
inelastic external chitinized cuticle, 
or exoskeleton. So at fixed periods 
(varying with the various species 
both in number and duration) this 
cuticle is cast or moulted. From 
a median longitudinal rent along 
the dorsum of the thorax and head, 
the insect, soft and dangerously 
helpless, struggles out of the old 
skin, enclosed in a new cuticle 
which, however, requires some little 
time to harden and assume its 
proper colors (often protective). 
After each moulting the young 
locust appears markedly larger and 
with its wing-pads better developed 
(Fig. 73). But not until the final 
moulting — in the case of the locust 
this is the fifth — are the wings usable as organs of flight. So that there 
is after all likely to be a rather marked difference between the habits of 
the young and those of the adult of an insect with incomplete metamor- 
phosis, that difference being primarily due to structural differences. The 
young are confined to the ground, and their locomotion is limited to walking 
or hopping. The adults can live, if they like, a life in the air, and they 
have a means of locomotion of greatly extended capability. 

The insects with complete metamorphosis are the beetles, the two- 
winged flies, the butterflies and moths, the ichneumons, gall-flies, ants, 
bees, and wasps, the fleas, the ant-lions, and several other small groups 
of insects with less familiar names. In the case of all the thousands of 
species in these groups, the young when hatched from the egg differ very 
much in structure and appearance, and also in habits and general economy, 
from the parents. Familiar examples of such young are the caterpillars 
and "worms" of the moths and butterflies, the grubs of beetles, the mag- 

FiG. 74. — Metamorphosis, incomplete, of an 
assassin-bug (family Reduviidae, order 
Hemiptera). A, young just hatching from 
eggs; B, young after first moulting, showing 
beginning wing-pads; C, older stage with 
complex wing-pads; D, adult with fully 
developed wings. (One-half larger than 
natural size.) 


Development and Metamorphosis 

gots of the flesh- and house-flies, and the helpless soft white grubs in the 
cells of bees and wasps. These strange young, so unlike their parents, 
have the generic name larva?, and the stage or life of the insect passed as a 
larva is known as the larval stage. In almost all cases these larvae have 
mouth-parts fitted for biting and chewing, while most of the adults have 
sucking-mouth parts; the larvae have only simple eyes and small inconspicu- 

FiG. 75. — Metamorphosis, complete, of monarch butterfly, Anosia plexippus. a, egg 
(greatly magnified); b, caterpillar or larva; c, chrysalid or pupa; d, adult or imago. 
(After Jordan and Kellogg. Natural size.) 

ous antennae; the adults have both simple and compound eyes and well- 
developed conspicuous antennae; the larvae may have no legs, or one pair or 
two or any number up to eight or ten pairs; the adults have always three 
pairs; the larvae are wholly wingless, nor do external wing-pads (i.e., 
developing wings) appear outside the body during the larval stage; the 
adults have usually two pairs (sometimes one or none) of fully developed 
wings. Internally the differences are also great. The musculation of the 

Development and Metamorphosis 


Fig. 76. — Larva, pupa, and adult of 

the flesh-fly, Calliphora crytJiroce- 
phala, with complete metamor- 
phosis. (Two times natural size.) 

larva is like that of a worm, to accomplish wriggling, crawling, worm-like 
locomotion; in the adult it is very different, particularly in head and thorax; 
the alimentary canal is usually adapted in the larva for manipulating and 
digesting solid foods; in the adult, usually (except with the beetles and 
a few other groups), for liquid food; there may be large silk-glands in the 
larva, which are rarely present in the 
adult; the respiratory system of the larvae 
of some flies and Neuroptera is adapted 
for breathing under water; this is only 
rarely true of the adults. The heart 
and the nervous system show lesser dif- 
ferences, but even here there is no iden- 
tity : the ventral nerve chain of the larvae 
may contain twice as many distinct gan- 
glia as in the adult. 

The larva lives its particular kind of 
life: it grows and moults several times; 
but externally it shows at no time any 
more likeness to the adult than it did at 
hatching. But after its last moult it ap- 
pears suddenly in the guise of a partially 
formed adult in (usually) quiescent mummy-like form, with the antennae, 
legs, and wings of the adult folded compactly on the under side of the 
body, and the only sign of life a feeble bending of the hind-body in re- 
sponse to the stimulus of a touch. This is the insect of complete meta- 
morphosis in its characteristic second stage (or third if the egg stage 

is called first), the pupal stage. The 
mummy is called pupa or chrysalid. As 
the insect cannot, in this stage, fight or 
run away from its enemies, its defence 
lies in the instinctive care with which the 
larva, just before pupation, has spun a 
protecting silken cocoon about itself, or 
has burrowed below the surface of the 
ground, or has concealed itself in crack 
or crevice. Or the defence may lie in the fine harmonizing of the color and 
pattern of the naked exposed chrysalid with the bark or twig on which it 
rests; it may be visible but indistinguishable. The insect as pupa takes 
no food; but the insect as larva has provided for this. By its greed and 
overeating it has laid up a reserve or food-store in the body which is drawn 
on during the pupal stage and carries the insect through these days or weeks 
or months of waiting for the final change, the transformation to the renewed 

Fig. 77. — Adult worker {a) and larva 
{b) of honey-bee. (Adult natural 
size; larva twice natural size.) 


Development and Metamorphosis 

active food-getting life of the adult or imaginal stage. Familiar examples 
of this kind of metamorphosis, the real metamorphosis, are provided by 
the life of the monarch butterfly, the honey-bee, and the blow-fly. The great 
red-brown monarch lays its eggs on the leaves of a milkweed; from the eggs 
hatch in four days the tiny tiger-caterpillars (larvae) (Fig. 75) with biting 
mouth-parts, simple eyes, short antennae, and eight pairs of legs on its elon- 
gate cylindrical wingless body. The caterpillars bite off and eat voraciously 
bits of milkweed-leaf; they grow rapidly, moult four times, and at the end 
of eleven days or longer hang themselves head downward from a stem or 

Fig. 78. — Brood-cells from honey-bee comb showing different stages in the metamor- 
phosis of the honey-bee; worker brood at top and three queen-cells below; begin- 
ning at right end of upper row of cells and going to left, note egg, young larva, old 
larva, pupa, and adult ready to issue; of the large curving queen-cells, two are cut 
open to show larva within. (After Benton; natural size.) 

leaf and pupate, i.e., moult again, appearing now not as caterpillars, but as 
the beautiful green chrysalids dotted with gold and black spots. The form- 
ing antennal legs and wings of the adult show faintly through the pupal 
cuticle, but motionless and mummy-like each chrysalid hangs for about 
twelve days, when through a rent in the cuticle issues the splendid butterfly 
with its coiled-up sucking proboscis, its compound eyes, long antennae, its 
three pairs of slender legs (the foremost pair rudimentary), and its four great 
red-brown wings. The queen honey-bee lays her eggs, one in each of the 
scores of hexagonal cells of the brood-comb (Fig. 78). From the egg there 
hatches in three days a tiny footless, helpless white grub, with biting mouth- 
parts and a pair of tiny simple eyes. The nurses come and feed this larva 
steadily for five days; then put a mass of food by it and "cap" the cell; the 
larva has grown by this time so as nearly to fill the cell. It uses up the 
stored food, and "changes" to the pupa, with the incomplete lineaments 
of the adult bee. It takes no more food, but lies like a sleeping prisoner 

Development and Metamorphosis 


in its closed cell for thirteen days, and then it awakens to active life, gnaws 
its way through the cell-cap and issues into the hive-space a definitive honey- 
bee with all the wonderful special structures that make the honey-bee body 
such an effective little insectean machine. The blow-fiy (Fig. 76) lays a hun- 
dred or more little white eggs on exposed meat. From these eggs come in 
twenty or thirty hours the tiny white wriggling larva: (maggots), footless, eye- 
less, wingless, nearly headless, with a single pair of curious extensile hooks 
for mouth-parts. For ten to fourteen days these larva; squirm and feed and 
grow, moulting twice in this time; they then pupate inside of the larval 
cuticle, which becomes thicker, firmer, and brown, so as to enclose the deli- 
cate pupa in a stout protective shell. The blow-fly now looks like a small 
thick spindle-shaped seed or bean, and this stage lasts for twelve or fourteen 

knitl. ■hm.l. h.p.1, 

Fig. 7q. — Dipterous larvae showing (through skin) the imaginal discs or buds of wings, 
these buds being just inside the skin. A, larva of black fly, Simulium sp.; B, anteiior 
end of larva of midge, Chironomus sp.; C, anterior end, cut open, of larva of giant 
crane-fly, Holorusia ruhiginosa;, bud of prothoracic respiratory tube;, 
bud of prothoracic leg;, bud of mesothoracic wing;, bud of mesothoracic 
leg; h.mtb., bud of metathoracic balancer; h.tntl., bud of metathoracic leg. (Much 

days. Then the winged imago, the buzzing blow-fly, as we best know it, 
breaks its way out. In the house-fly the same kind of life-history, with 
complete metamorphosis of the extremest type, is completed in ten days. 
Nor do we realize how really extreme and extraordinary this metamorpho- 
sis is until we study the changes which take place inside the body, as well 
as those superficial ones we have already noted. 

The natural question occurs to the thoughtful reader: "Is the meta- 
morphosis or transformation in the postembryonal development of such 
insects as the butterfly, bee, and blow-fly as sudden or discontinuous and 
as radical as the superficial phenomena indicate? " The answer is no, and 
yes; the metamorphosis is not so discontinuous or saltatory and yet is 
even more radical and fundamental than the external changes suggest. To 


Development and Metamorphosis 

take a single example, the case of the blow-fly (admittedly an extreme one), 
the phenomena of internal change are, put briefly, as follows: The imaginal 
wings, legs, and head-parts begin to develop as deeply invaginated httle 
buds of the cell-layer of the larval skin early in larval life. This develop- 
ment is gradual and continuous until pupation, when the wing and le^^ rudi- 

FiG. 80. — Stages in development of wing-buds in the larva of the giant crane-fly, 
Holorusia riibiginosa (the wing-buds have been dissected out and sectioned, so 
as to show their intimate anatomy). A, B, C, D, four stages successively older ch., 
chitinized cuticle; hyp., hypoderm or cellular layer of skin; tr., trachea; irl., 
tracheoles; p.m., peritrophic membrane; w., developing wing; t.v., tracheal branch 
indicating position of future wing-vein. (Greatly magnified.) 

ments and the new head are pulled out upon the exterior of the body. Just 
before pupation, when the larva has given up its locomotion and feeding, 
the larval muscles, tracheae, salivary glands, alimentary canal, and some other 
tissues begin to disintegrate, and rapidly break wholly down, so that in the 
pupa there appear to be no internal organs except the nervous system, 
reproductive glands, and perhaps the heart, but the whole interior of the 

Development and Metamorphosis 





body is filled with a thick fluid in which float bits of degenerating larval 
tissue. At the same time with this radical histolysis or breaking down of 
tissue a rapid histogenesis or developing of imaginal parts from certain 
groups of undifferentiated primitive cells, derived probably mostly from 
the larval skin-cells, is going on. Thus many of the larval organs and tissues, 
instead of going over into the corresponding imaginal ones, wholly disinte- 
grate and disappear, and the imaginal parts are newly and independently 
derived. In connection with the 
breaking down of the larval tissues 
phagocytes or freely moving, tissue- 
eating, amoeboid blood-cells play an 
important part, although one not 
yet fully understood. They are 
either the causal agents of the 
histolysis, or are assisting agents in 
it, the tissue disintegration beginning 
independently, or — a recent sugges- 
tion — they are perhaps more truly 
to be looked on as trophocytes, 
that is, carriers of food, namely, 
disintegrating tissue, to the develop- 
ing centers of the imaginal parts. 
Much investigation remains to be 
done on this interesting subject 
of histolysis and histogenesis in 
insects with complete metamor- 
phosis, but enough has been already accomplished to show the basic and 
extreme character of the transformation from larva to adult. 

If we ask for the meaning of such unusual and radical changes in the 
development of insects, we confront at once an important biological prob- 
lem. Most biologists believe that in a large and general way the develop- 
ment of animals is a swift and condensed recapitulation of their evolution; 
meaning by development the life-history or ontogeny of an individual, and 
by evolution the ancestral history or phylogeny of the species. According 
to this "biogenetic law" the interpretation of the significance of the various 
stages and characters assumed by an animal in the course of its development 
from single fertilized egg-cell to the complex many-celled definitive adult 
stage is simple: These stages correspond to various ancestral ones in the 
long genealogical history of the species. Every vertebrate, for example, is 
at some period in its development more like a fish than any other living 
kind of animal ; it has gill-slits in its throat, is tailed, and is indeed a fish- 
like creature. This is its particular developmental stage, corresponding 

Fig. 8i. — A cross section of the body of the 
pupa of a honey-bee, showing the body-cavity 
filled with disintegrated tissues and phago- 
cytes, and (at the bottom) a budding pair 
of legs of the adult, the larvK being 
wholly legless. Photomicrograph by George 
O. Mitchell; greatly magnified.) 


Development and Metamorphosis 

to the ancestral fish-like ancestors of all vertebrates. Do then the larvae 
and pupae of insects with complete metamorphosis represent ancestral stages 
in insect evolutionary history? In some degree the larval stage does, but 

in no degree does the pupal. 
Insects are certainly not de- 
scended from an animal that, 
like a pupa, could neither move 
nor eat and which had no in- 
ternal organs except a nervous 
system, heart, and rudimentary 
reproductive glands. Biologists 
recognize that the exigencies of life during adolescence may profoundly 
modify what might be termed the normal course of development. As 
long as the developing animal is shielded from the struggle for existence, 
is provided with a store of food and protected from enemies by lying in an 
egg-shell or in the body of the mother, it may pursue fairly steadily its reca- 
pitulatory course of development; but once emerged and forced to shift for 

Fig. 82. — A bit of degenerate muscle from tussock 
moth, Hemerocampa leucostigma. Note phago 
cytic cells attacking muscle at the margins 
(Greatly magnified.) 

Fig. 83. — Degenerating muscle from pupa of giant crane-fly, Holorusia ruhiginosa, show- 
ing phagocytic cells penetrating and disintegrating the muscle -tissue. (Greatly 

itself, it must be, at whatever tender age it is turned out, or whatever ancient 
ancestor it is in stage of simulating, adapted to live successfully under the 
present-day and immediate conditions of life. If the butterfly gets hatched 
long before it has reached its definitive butterfly stage, and while it is in 
a stage roughly corresponding to some worm-like ancestors — and from such 
ancestors insects have undoubtedly descended — it must be fitted to live 

Development and Metamorphosis 


successfully a crawling, squirming, worm-like life. That those insects which 
hatch as worm-like larvae do in fact owe their wingless, worm-like body con- 
dition partly to being born in a stage simulating a worm-like ancestor is proba- 

FiG. 84. — Degeneration, without phagocytosis, of salivary glands in old larva of giant 
crane-fly, Holorusia rubiginosa. A, cross-section of salivary gland before degen- 
eration has begun; B, cross-section of salivary gland after degeneration has set in. 
(Greatly magnified.) 

bly true. But to be a successful worm demands very different bodily adapta- 
tions from those of a successful butterfly. And so far does the larval butterfly 
go, or so far has it been carried, in meeting these demands that nature finds it 
more economical — to get into figurative language — 
or easier to break down almost wholly the larval 
body — after a new food-supply for further develop- 
ment has been got and stored away, and to 
build up from primitive undifferentiated cell begin- 
nings the final definitive butterfly body, than to 
make over these very unlike larval parts into the 
adult ones. The pupal stage, quiescent, non-food 
taking, and defended by a thick chitinous wall, 
often enclosed in a silken cocoon, buried in the 
ground or crevice, or harmonizing so perfectly with 
its environment as to be indistinguishable from it, 
is the chief period of this radical and marvelous 
breaking down and building anew. It is an inter- 
polated stage in the development of the butterfly 
corresponding to nothing in the phyletic history; 
an adaptation to meet the necessities of its life- 
conditions. To my mind, this is the interpretation of the phenomena of 
complete metamorphosis. 

Fig. 85. — Cross-section 
of newly developing 
muscle in pupa of 
honey-bee, Apis mel- 
lifica. (Greatly mag- 


As has been explained in the preceding chapter, insects are primarily classi- 
fied on the basis of their postembryonic development. Insects with incom- 
plete metamorphosis, that is, those which do not undergo a non-feeding, 
usually quiescent, pupal stage in their development are believed to be more 
nearly related to each other than to any of the insects which undergo a so- 
called complete metamorphosis. So they are spoken of collectively as the 
Hemimetabola, while all the insects with a distinct pupal stage are called 
the Holometabola. But when one has collected an adult insect, as a fly 
or moth or grasshopper, and wishes to classify it, this primary classification 
based on character' of development often cannot be made for lack of informa- 
tion regarding the life-history of the particular insect in hand. The next 
grouping is into orders, and this grouping is based chiefly on structural 
characters, and corresponds to one's already more or less familiar knowledge 
of insect classification. Thus all the beetles with their horny fore wings 
constitute one order, the Coleoptera; the moths and butterflies with their 
scale-covered wings another order, the Lepidoptera; the two-winged flies 
the order Diptera, the ants, bees, wasps, and four-winged parasitic flies 
the order Hymenoptera, and so on. So that the first step in a beginner's 
attempt to classify his collected insects is to refer them to their proper orders. 

Now while entomologists are mostly agreed with regard to the make-up 
of the larger and best represented orders, that is, those orders containing 
the more abundant and familiar insects, there are certain usually small, 
obscure, strangely formed and more or less imperfectly known insects with 
regard to whose ordinal classification the agreement is not so uniform. While 
some entomologists incline to look on them simply as modified and aberrant 
members of the various large and familiar orders, others prefer to indicate 
the structural differences and the classific importance of these differences 
by establishing new orders for each of these small aberrant groups. IVIost 
entomologists of the present incline toward this latter position, so that whereas 
Linnasus, the first great classifier of animals, divided all insects into but 
seven orders, the principal modern American * text-book of systematic ento- 

* Comstock, J. H., A Manual of Insects, 1898. 


The Classilication of Insects 53 

mology recognizes nineteen distinct ones. This does not mean, of course, that 
twelve new orders of insects have been found since Linnaius's time, although 
two or three of the orders are in fact founded on insects unknown to him, 
but means that certain small groups classified by Linnaeus simply as famihes 
in his large orders have been given the rank of distinct orders by modern 
systematists. And as our knowledge of insects and their relationship to 
each other is certainly much larger now than it was one hundred and fifty 
years ago, we may feel confident that the many-order system of classifica- 
tion is more nearly a true expression of the natural interrelationships of 
insects than was the old seven-order system. But not all entomologists 
agree on the nineteen-order system. Few, indeed, still use the Linnaean 
system, but many believe that the division of the insect class into nineteen 
orders gives too much importance to certain very small groups and to some 
others which are not markedly aberrant, and these entomologists recognize 
a lesser number of orders, varying with different authors from nine to about 
a dozen. In this book we shall adopt the nineteen-order system as used 
in Comstock's Manual. In the first place the author beheves that this classi- 
fication best represents our present knowledge of insect taxonomy; in the 
second place this is the classification taught by nearly all the teachers of 
entomology in America. 

To determine the order to which an insect belongs we make use of a 
classifying table or key. In the Key to Orders which follows this para- 
graph, all the insect orders are characterized by means of brief statements of 
structural features more or less readily recognized by simple inspection of 
the superficies of the body; to determine some of the conditions a simple 
lens or hand-magnifier will be needed. The orders are so arranged in the 
key that by choosing among two or more contrasting statements the student 
may "trace" his specimen to its proper order. Inspection of the Key with 
an attempt or two at tracing some familiar insect, as a house-fly, moth, or 
wasp whose order is already known, will make the method of use apparent. 
It must be borne in mind that young insects, such as caterpillars of moths, 
grubs of beetles, and the wingless nymphs of locusts, dragon-flies, etc., cannot 
be classified by this key. Indeed the young stages of most of the insects 
which we know well as adults are unknown to us, and there is, besides, such 
manifold adaptive variety in the external structure of those forms which we 
do know that no key for the classification into orders of immature insects 
can now be made. 

54 The Classification of Insects 

(Arranged by Prof. H. E. Summers.) 

(For adult insects only. If in any paragraph all the italicized characters agree with 
the specimen in hand, the remaining characters need not be read; these latter are for use 
in doubtful cases, or where the organs characterized in italics are rudimentary or absent. 
The technical terms used in this Key have all been defined in Chapter I.) 

A. Primitive wingless insects; month-parts well developed, hut all except the apices oj the 
mandibles and maxillcB withdrawn into a cavity in the head; tarsi (feet) always one- 
or two-clawed; body sometimes centiped-like, with well-developed abdominal legs, 

in this case tarsi two-clawed (The simplest insects.) Aptera. 

AA. Normally winged insects, wings sometimes rudimentary or absent; mouth -parts 
not withdrawn into a cavity in the head. 

B. Month-parts, when developed, with both mandibles and maxillce fitted for biting; 
abdomen broadly joined to thorax; tarsi never bladder-shaped; when mouth- 
parts are rudimentary, if the wings are two, there are no halteres (p. 303) ; if 
the wings are four or absent, the body is not densely clothed with scales. 
C. Posterior end of abdomen with a pair of prominent tmjointed forceps-like 
appendages; fore wings, when present, short, veinless, horny or leathery. 

(Earwigs.) Euplexoptera. 
CC. Posterior end of abdomen usually without prominent unjointed forceps-like 
appendages; when these are present the fore wings are always developed, 

D. Fore wings, when present, veined and membranous, parchment-like or 
leathery; when absent, the labium (under-lip) either cleft in the 
middle, or the mouth-parts prolonged into a distinct beak. 
E. Fore wings, when present, thicker than hind wings, somewhat 
leathery or parchment-like; hind wings folded several times 
lengthwise, like a fan, in repose; when wings are absent, pro- 
thorax large. 

(Locusts, crickets, cockroaches, etc.) Orthoptera. 
EE, Fore wings membranous, of same structure as hind wings; 
hind wings usually not folded, but occasionally folded like a fan; 
when wings are absent, prothorax small. 
F, Antennce inconspicuous. 

G. Hind wings smaller than fore or absent; posterior end of 

abdomen with two or three many-jointed filaments. 

(May -flies.) Ephemerida. 

GG. Hind wings not smaller than fore; posteiior end of 

abdomen without many-jointed filaments. 

(Dragon-flies and damsel-flies.) Odonata. 
FF. Antennce conspicuous. 

G. Tarsi less than five-jointed; labium cleft in the 

H. Wings always present, although sometimes very 
small; hind wings broader than fore wings, 
folded in repose; prothorax large, nearly flat 
on dorsal surface. 

(Stone-flies.) Plecoptera. 

The Classification of Insects ^^ 

HH. Hind wings, when present, not broader than fore 
wings, not jolded in repose; protliorax small, 

I. Tarsi four-jointed; wings, when present, 
equal in size (Termites.) Isoptera. 

II. Tarsi one- to three-jointed. 

J. Tarsi one- or two-jointed; ahvays 

(Biting bird-lice.) Mallophaga. 

JJ. Tarsi usually three-jointed ; occasionally 

two-jointed, in which case wings always 

present, fore wings larger than hind 

wings. (Book -lice, etc.) Corrodentia. 

GG. Tarsi five-jointed, but with one joint sometimes 

difficult to distinguish; labium usually entire in 

middle, sometimes slightly emarginate. 

H. Wings, when present, naked or slightly hairy; 

hind wings with or without jolded anal space; 

in former case prothorax large and nearly 

flat on dorsal surface; in wingless forms 

mouth prolonged into a distinct beak. 

I. Mouth-parts not prolonged into a distinct 
beak, at most slightly conical. 

(Dobsons, ant-lions, etc.) Neuroptera. 

II. Mouth-parts prolonged into a distinct beak. 

(Scorpion-flies, etc.) Mecoptera. 
HH. Wings, when present, thickly covered with hairs; 
hind wings usually with jolded anal space; pro- 
thorax small, collar-like; mouth not prolonged 
into a beak. (Caddis-flies.) Trichoptera. 
DD. Fore wings, when present, veinless; horny or leathery; when absent, 
labium entire, and mouth-parts not prolonged into a distinct beak. 

(Beetles.) Coleoptera. 
BB. Mouth-parts, when developed, more or less fitted for sucking; sometimes also 
fitted in part (the mandibles) for biting: in this case either (i) base of abdomen 
usually strongly constricted, joined to thorax by a narrow peduncle, or (2) the 
tarsi bladder -shaped, without claws; when mouth is rudimentary either the 
wings are two and halteres are present, or the wings are four or none and 
the body (and wings if present) are densely clothed with scales. 
C. Prothorax jree; body {and wings if present) never densely clothed with 
scales; maxillary palpi usually absent; when present, tarsi bladder- 
shaped, without claws. 

D. Tarsi bladder-shaped, without claws; wings four {sometimes absent), 
narrow, fringed with long hairs; maxillae triangular, with palpi. 

(Thrips.) Thysanoptera. 

DD Tarsi not bladder-shaped, usually clawed; wings not fringed with 

long hairs; maxilla {when mouth is developed) bristle-like, without 

palpi. (Bugs.) Hemiptera. 

CC. Prothorax not free; maxillary palpi present, sometimes rudimentary 

and difficult to see, in which case body (and wings if present) densely 

clothed with scales; tarsi never bladder-shaped, usually clawed. 

56 The Classification of Insects 

D. Mandibles often rudimentary, when present bristle-like. 

E. Wings jour {sometimes wanting), clothed with scales; body 
covered thickly with scales or hairs; mouth, when developed, a 
slender sucking proboscis, closely coiled under head. 

(Moths and butterflies.) Lepidoptera. 
EE. Wings two {or wanting), naked or with scattered hairs; hind 
wings in winged forms represented by halteres; body either 
naked or with scattering hairs; mouth a soft or horny beak, not 
coiled under head. 
F. Prothorax poorly developed, scarcely visible from dorsal 

side (Flies.) Diptera. 

FF. Prothorax well developed, distinctly visible from dorsal 

side; wings never present (Fleas.) Siphonaptera. 

DD. Mandibles well developed, fitted for biting; wings four {sometimes 
two or none), naked or with scattered hairs. 
(Ichneumon-flies, gall-flies, wasps, bees, and ants.) Hymenoptera. 

After one has classified an insect in its proper order there remains, first, 
the determination of the family (each order being composed of from one 
to many families), then of the genus (each family comprising one to many 
genera), and finally of the particular species of the genus (each genus includ- 
ing one to many species). This ultimate classification to species, however, 
will be possible to the amateur in comparatively few cases. There are so 
many species of insects (about 300,000 are known) that it would require 
many shelves of books to contain the descriptions of them all. As a matter 
of fact, in only a few orders have the descriptions of the species been brought 
together in manuals available for general students. For the most part the 
descriptions are scattered in scientific journals printed in various languages 
and wholly inaccessible to the amateur. There are less than 1000 different 
species of birds in North America; there are more than 10,000 known 
species of beetles. Now when one recalls the size of the systematic man- 
uals of North American birds, and realizes that ten such volumes would 
include only the insects of one order, it is apparent that complete manuals 
of North American insects are out of the question. Except in the case of 
the most familiar, wide-spread, and readily recognizable insect species we 
must content ourselves with learning the genus, or the family, or with the 
more obscure, slightly marked, and difficult members of certain large groups, 
as the beetles and moths, simply the order of our insect specimens. 

When one has determined the order of an insect by means of the above 
key he should turn to the account of this particular order in the book (see 
index for page) and find the keys and aids to the further classification of 
the specimen which the author has thought could be used by the general 
student. Comparison with the figures and brief descriptions of particular 
species which are given in each order may enable the amateur to identify 
the exact species of some of his specimens. But the specific determination 

The Classification of Insects ^J 

of most of the insects in an amateur's cabinet (or in a professional ento- 
mologist's either, for that matter) will have to be done by systematic 
speciahsts in the various insect groups. Few professional entomologists 
undertake to classify their specimens to species in more than the one or 
two orders which they make their special study. Duplicate specimens should 
be given numbers corresponding to those on specimens kept in the cabinet, 
and be sent to specialists for naming. Such specialists, whose names can 
be learned from any professional entomologist, have the privilege of retain- 
ing for their own collections any of the specimens sent them. 


ERTAIN household pests which are 
not moths and do not look like 
fish, but which are com ' ^^^_Jg monly called "fish-moths" (Fig. 86), are 
our most familiar repre sentatives of the order of "simplest in- 

sects." The "fish" part of the name comes from the 
covering of minute scales which gives the body a silvery 
appearance, and the "moth" part is derived from our 
habit of calling most household insect pests "moths." 
Thus we speak of "buffalo-moths" when we refer to the 
carpet-feeding hairy larvae of certain beetles. When we 
say clothes-moths we are really using the word moth 
accurately, for in their adult condition these pests are 
true moths, although the injury to clothing is wholly done 
by the moth in its young or caterpillar stage. 

Besides the fish-moths other not unfamiliar Aptera are 
the tiny "springtails" (Fig. 87), which sometimes occur 
in large numbers on the surface of pools of water or on 
snow in the spring. Others may be easily found in damp 
decaying vegetable matter, as discarded straw or old toadstools. They are 
provided with an odd little spring on the under side of the body by means 
of which they can leap from a few inches to a foot 
or more into the air. Hence their common name. 

In the order Aptera are included the simplest of 
living insects. By "simplest" is meant most primi- 
tive, most nearly related to the ancestors of the whole 
insect class. Also, as might be expected, these most 
primitive insects are simplest in point of bodily struc- 
ture; but in this respect they are nearly approached 
by simple-bodied members of several other orders. 
These latter forms, however, have a simple body- 
structure due to the degradation or degeneration of a more complex type. 


Fig. 86.— The fish- 
moth, Lepisma 
saccharina. (After 
Howard and Mar- 
latt; twice natural 

Fig. 87. — The pond-sur- 
face springtail, Smyn- 
thurus aquations. 
(After Schott; much 

The Simplest Insects 


Fig. 88. — Diagrammatic figures 
showing the segmental disposi- 
tion of the ovarial tubes in three 
Apteran genera. A, J a pyx; B, 
Lepisma; C, Campodea. (After 
Targioni-Tozzetti; much en- 

It is familiar knowledge that animals which live parasitically on others, or 

which adopt a very sedentary life, show a marked degeneration of body 

structure, an acquired simplicity due to the loss of certain parts, such as 

organs of locomotion (wings, legs), and of 

orientation (eyes, ears, feelers, etc.). Thus 

the parasitic biting bird-lice (order Mal- 

lophaga, see p. 113), which Uve their whole 

lives through on the bodies of birds, feeding 

on the feathers, are all wingless and of gener- 
ally simple superficial structure. They are 

nearly as simple externally perhaps as the 

Aptera, but we believe that they are the 

degenerate descendants of winged and in 

other ways more complexly formed ancestors. 
Similarly certain species of insects in 

nearly all orders have adopted a life-habit 

which renders flight unnecessary, and these 

insects having lost their wings are in this 

character simpler than the winged kinds. 

Examples of such insects are the worker 

ants and worker termites, many household insects, as the bedbugs and fleas, 

and many ground-haunting forms, as some 
of the crickets, cockroaches, and beetles. 

The Aptera, however, owe their sim- 
plicity to genuine primitiveness; among all 
living insects they are the nearest repre- 
sentatives of the insectean ancestors. But 
not all the Aptera are "simplest." That 
is, within the limits of this small order a 
considerable complexity or specialization of 
structure is attained, although all the 
Aptera are primitively wingless, as the 
name of the order indicates. 

These insects develop "without meta- 
morphosis"; that is, the young (Figs. 90 
and 94) are almost exactly like the parents 

Fig. 89.— Diagrammatic figures show- except in size. They have simply to grow 
ing the respiratory system in three larger and to become mature. In internal 
Apteran genera. A, Machilis; B, ^ ^ ^1 • 1 a .l 1 

Nicoletia; C, Jcipyx. (After Tar- Structure the smipler Aptera show some 
gioni-Tozzetti; much enlarged.) most interesting conditions. Their internal 

systems of organs have a segmental character corresponding to the external 

segmentation of the body. The ovarial tubes, which are gathered into 

6o The Simplest Insects 

two groups or masses, one on each side of the body, in all other insects 
(Fig. 66), are separate and arranged segmentally in Japyx (Fig. 88), and 
less markedly so in Machilis; the respiratory system of Machilis (Fig. 89) 
consists of nine pairs of distinct, segmentally arranged groups of tracheae 
(air-tubes), while the ventral nerve-cord has a ganglion in almost every seg- 
ment of the body. As insects are certainly descended from ancestors whose 
bodies were composed of segments much less interdependent and coordi- 
nated than those of the average living insect, those present-day insects which 
have the body both externally and internally most strongly segmented are 
believed to be the most generalized or primitive of living forms. In addi- 
tion to the segmented character of the internal organs we have also another 
strong evidence of the primitiveness of the order in the possession by several 
Aptera of rudimentary but distinct external pairs of appendages on the 
abdominal segments, appendages undoubtedly homologous with the thoracic 
legs, and probably well developed in the insect ancestors as abdominal legs 
like those of the centipeds. 

The order Aptera is composed of two suborders, which may be dis- 
tinguished as follows: 

Abdomen elongate, composed of ten segments, and bearing long bristle-like or 
shorter forceps-like appendages at its tip; no sucker on ventral side of first 
abdominal segment; antennae many-segmented Thysanura. 

Abdomen short and robust, composed of six segments, and usually with a forked 
spring at tip (usually folded underneath the body), and with a ventral sucker 
on first abdominal segment; antennae 4- to 8-segmented Collembola. 

Th\'Sanura. — This suborder includes three families (a problematical 
fourth family is found in Europe), as follows: 

Body covered with scales Lepismid.e 

Body not covered with scales. 

Tip of abdomen with forceps-like appendages jAPYGiDiE. 

Tip of abdomen with slender many-segmented appendages Campodeid^. 

To the last family in the above key belongs the interesting creature 
Catnpodea staphylinns (Fig. 90), which zoologists regard as the most primi- 
tive living insect. It is small, white, flattened, wingless, and so soft-bodied 
and delicate that it can hardly be picked up uninjured with the most deli- 
cate forceps. It is about \ inch long (exclusive of caudal appendages), and 
is to be looked for under stones and bits of wood. I have found it in Ger- 
many, in New York, and in California, which indicates its wide distribu- 
tion. Other collectors have taken it in Italy, England, and in the Pyrenees. 
It is said to live also in East India. Is it not a little surprising that this 
most primitive, wholly defenceless, and ancient insect should be able to live 
successfully the world over in the face of, and presumably in competition 
with, thousands of highly developed specialized modern insect forms? It 

The Simplest Insects 


is a striking proof that Nature does not inevitably crush out all of her 
first trials in favor of her later results! 

The Campodeidte contain another 
genus, Nicoletia (Fig. 91), one species of 
which, A^. iexensis, has been found in Cali- 
fornia and Texas, and which may be dis- 
tinguished from Campodea by its posses- 
sion of three caudal appendages instead 
of two as in the latter form. 

The Japygidae include but a single 
genus, Japyx, represented in this country 
by two described species and several as yet 
undescribed forms found at Stanford Uni- 
versity. Japyx siihterraneus is a species 
first found under stones at the mouth of 
a small grotto near the Mammoth Cave 
(Kentucky). Japyx (Fig. 92) is larger Fig. go 
than Campodea,^being about one-half inch 
long, and is readily recognized by its caudal 

Young and adult of Cam- 
podea slaphylinus (from California), 
the simplest living insect. (Natural 
size indicated by line.) 

forceps. Like Campodea its body is white and soft. 

The Lepismidae include the familiar household fish- 
moths and a number of similar forms which live under 
stones and logs in soft soil at the bases of tree-trunks, 
under dead leaves in woods, and sometimes on the damp 
sand of seashores. Three genera of this family occur 
in North America, which may be distinguished as 

Caudal appendages short; prothorax very wide and body 

behind it tapering rapidly Lepismina. 

Caudal appendages long; body elongate and tapering 
gradually backward. 

Eyes large and close together Machilis. 

Eyes small and far apart Lepisma. 

Lepisma is best known by the species L. saccharina 
(Fig. 86), which is the silverfish or fish-moth of the 
house. It is silvery white, with a yellowish tinge on 
the antennae and legs, and is from one-third to two- 

p V Z /■ c fifths of an inch long. The three long caudal appen- 

ensis, from Califor- dages, characteristic of the genus, are conspicuous. It 

ma. (Eight times nat- fg^^jg chiefly on sweet or starchy materials, sometimes 

doing much damage in libraries, where it attacks the 

bindings. It attacks starched clothing, eats the paste off the wall-paper, 


The Simplest Insects 

causing it to loosen, and infests dry starchy foods. It runs swiftly and 
avoids the light. It can be fought by sprinkling fresh 
pyrethrum powder in bookcases, wardrobes, and 
pantries. Another species, L. domestica (Fig. 93), 
called the bake-house silverfish, is often common 
about fireplaces and ovens, running over the hot 
metal and bricks with surprising immunity from the 
effects of the heat. This habit has gained for it in 
England, according to Marlatt, the name of "fire- 
brat." It can be distinguished from the species 
saccharina by the presence of dark markings on the 

Fig. 92. — Japyx sp., from back. Both saccharina and domestica are common 
California. (Five times jj^ England, and saccharina probably came to this 
natural size.) r 1 

country from there. 

Machilis (Fig. 95) does not occur in houses, but is more common than 
Lepisma outdoors. It is to be found under stones, in the soil around the 
base of tree-trunks, among dead leaves and fallen pine-needles, and at least 
one species occurs in the sand of sea-beaches. 

Fig. 93. 

Fig. 94. 

Fig. 93. — The fish-moth, Lepisma domestica. (After Howard and Marlatt; a little 

larger than natural size.) 
Fig. 94. — Young and adult of Lepisma sp., from California. (Twice natural size.) 

CoLLEMBOLA. — The springtails, mostly of microscopic size, and wholly 
unfamiliar to any but persistent explorers of nature, comprise many more 
species than the Thysanura. Their most distinctive character is the pos- 
session, by most of them, of the forked spring (Figs. 96 and 97), by 
means of which they leap vigorously when disturbed. This spring is 

The Simplest Insects 


attached to the next to last body segment or to the antepenultimate one. 
It consists of a basal part and of two terminal processes. 
It is carried bent forward under the body, with the bipartite 
tip held in a little catch on the third abdominal segment. 
In some species the catch is lacking. The springtails also 
possess a curious organ on the ventral aspect of the first 
abdominal segment which appears to be a small projecting 
sucker or tube. This sucker is often more or less divided 
into two parts, in one family consisting plainly of two 
elongate, delicate tubes (Figs. 96 and 97). The use of 
this peculiar structure has not been definitely determined. 
Some entomologists think that it serves as a clinging organ, 
enabhng the insect to attach its body firmly to the object 
upon which it rests. Others believe that the sucker serves 
in some way to take up moisture, while still others be- 
lieve it to aid in respiration. The Collembola as well 
as the Thysanura cannot live in a dry atmosphere. 
This suborder is divided into five families, as follows 
(MacGillivray) : 

Fig. 95. — Machi- 
lis sp., from Cali- 
fornia. (Three 
times natural 

A. Spring wanting Aphorurid.i;. 

AA. Spring present. 

B. Spring arising from ventral side of 
antepenultimate abdominal segment. 


BB. Spring arising from ventral side of penultimate abdom- 
inal segment. 
C. Abdomen elongate, cylindrical, much longer than 

broad Entomobryid^. 

CC. Abdomen globular, but little larger than broad. 

D. Terminal segment of antennas long, ringed. 


DD. Terminal segment of the antennae short, with 

* a whorl of hairs Papiriid^. 

Fig. 96. — The spotted 
springtail, Papirius 
7naci(losus,with spring 
folded underneath 
body. (Natural 
length, 2 mm.) 

Of these five families the members of one, the Aphorurida?, in which 
the spring is wanting, are non-saltatorial. In all of 
the others leaping is a characteristic habit. The 
Smynthuridae and the Papiriidae are represented by 
but one genus each, viz., Smynthurus and Papirius. 
Smynthurus hortensis is a common form in gardens, 
and may be called the "garden-flea." It is found 
in the Eastern States in May and June "upon the Fig. 97.— The spotted 
leaves of young cabbage, turnip, cucumber, and £™,^;^i|; 3%'Se3ei 
various other plants, and also on the ground. It (Natural length, 2 mm.) 


The Simplest Insects 

is dull black, with head, legs, and bases of the antennae rust-color." Smyn- 
thurus aquaticus (Fig. 87) often occurs in great numbers on the surface of 
pools. The insects look like tiny black spots on the water surface, but a 

little observation soon reveals their 
lively character. 

The Poduridae and Entomobryidae 

are represented in North America by 

twelve and fourteen genera respec- 

>^am: tively. Many of the Podurids are 

illf I f \w\Vi||'j|iM >^^H^ covered with scales and are often 

mWmhJm ,^SSsL prettily colored and patterned. The 

scales (Fig. 98) are very minute and 
bear many fine lines and cross-lines, 
regularly arranged. On this account 
Fig. 98. Fig. 99. they are much used as test objects 

Fig. 98.— Scales from a springtall. (After ^^^ microscopes, the quality of the 

Murray; greatly magnified.) '■ • , , • 

Fig. 99.-^The snov^-fita, Achorutes nivicola. lens being determined by its capacity 
(After Folsom; much enlarged.) ^-q reveal their extremely fine mark- 

ings. One of the most interesting Podurids is 
the snow-flea, A chorutes nivkola (Fig. 99) , which 
gathers in large numbers on the surface of snow 
in the late spring. Comstock says that the 
snow-flea is sometimes a pest where maple- 
sugar is made, the insects collecting in large 
quantities in the sap. 

An interesting representative of the Entomo- 
bryidae is the house springtail, Lepidocyrtiis anieri- 
canus (Fig. 100), said by Marlatt to be "not 
infrequently found in dweUings in Washington." 
It is about one-tenth of an inch long, silvery 
gray, with purple or violet markings. In Europe 
also one species of springtail is common in 
houses. As these insects live on decaying vege- 
table matter, they probably do no special harm 
in the house. They especially frequent rather moist places, and may often 
be found in window-plant boxes and conservatories. 

Fig. 100. — The American 
springtail, Lepidocyrtus 
americanus, ventral aspect, 
showing spring folded un- 
derneath body. (After 
Howard and Marlatt ; 
much enlarged.) 


THE MAY-FLIES (Order Ephemerida) and STONE- 
' ' — FLIES (Order Plecoptera) 

AY-FLIES, lake-flies, or shad-flies, common names for 
the insects of the order Ephemerida, are famihar to 
people who live on the shores of lakes or large rivers, 
but are among the unknown insects to most high-and- 
dry dwellers. 

Travelling down the St. Lawrence River from 
Lake Ontario to Quebec one summer, I had hosts of 
day-long companions in little May-flies that clung to 
my clothing or walked totteringly across my open book. The summer 
residents of the Thousand Islands get tired of this too-constant com- 
panionship, and look resentfully on the feeble shad-fly as an insect pest. 
One evening in August, 1897, my attention, with that of other strollers along 
the shore promenade at Lucerne, was called to a dense, whirling, tossing 
haze about a large arc light suspended in front of the great Schweizerhof. 
Scores of thousands of May-flies, just issued from the still lake, were in 
violent circling flight about the blinding light, while other thousands were 
steadily dropping, dying or dead, from the dancing swarm to the ground. 
Similar sights are familiar in summer-time in this country about the lights 
of bridges, or lake piers and shore roads. This flying dance is the most 
conspicuous event in the life of the fully developed, winged May-fly, and 
indeed makes up nearly all of it. With most species of May-flies the winged 
adult lives but a few hours. In the early twilight the young May-fly floats 
from the bottom of the lake to the surface, or crawls up on the bank, the 
skin splits, the fly comes forth full-fledged, joins its thousands of issuing 
companions, whirls and dances, mates, drops its masses of eggs on to the 
the lake's surface, and soon flutters and falls after the eggs. It takes no 
food, and dies without seeing a sunrise. Sometimes the winds carry dense 
clouds of May-flies inland, and their bodies are scattered through the streets 
of lakeside villages, or in the fields and woods. Sometimes the great swarms 



The May-flies and Stone-flies 

fall to 


the water's surface and there are swept along by wind and wave, 
until finally cast up in thick winrows, miles long, 
on the lake beach. Millions of dead May-flies 
are thus piled up on the shores of the Great 

We call the May-flies the Ephemerida, after 
the Ephemerides of Grecian mythology, and the 
name truly expresses their brief existence — above 
water. But they have lived for a year at least 
before this, or for two or even three years, as 
wingless, aquatic creatures, clinging concealed 
to the under side of stones in the lake or stream 
bottom, or actively crawling about after their food, 
which consists of minute aquatic plants and animals 
or bits of dead organic matter. In this stage their 
whole environment, habits, and general appearance are 
radically different from those of the brief adult life. We 
can only guess, if our curiosity compels us to attempt some 
explanation, at the manner and the cause of such a 
strange life-history. What advantage is there in such a 
specialized condition that Nature could not have arrived 
at by less indirect means? What is indeed the utility of 
the whole modification? The quick answer "utility," 
which is to account for all such strange structural and 
physiological conditions on the basis of useful adapta- 
tions brought about by the slow but persistent action 
of natural selection, leaves us, confessedly, answered 
simply on a basis of belief. In hundreds of cases that 
may come under our observation, in how few are we 
really able to perceive a reason-satisfying course of adap- 
tive development based on the selection of useful small 
fluctuating variations? 

The eggs of the May-fly fall from the body of the 

mother to the water's surface in two packets, which, 

If however, break up while sinking, so that the released 

Fig. ioi. — May-flies about an electric lamp. 

The May-flies and Stone-flies 


eggs reach the bottom separately. From each egg hatches soon a thiy 
flattened, soft-bodied, six-legged creature called a nymph, without wings 
or wing-pads, and looking very much like a Campodea (the simplest 
living insect, see p. 61). This nymph crawls about, feeds, grows, moults, 
grows, moults again and again (in a species observed by Lubbock there 
were twenty-one moultings), and finally at the end of a year, or of two or 
three years, depending on the species, is ready to issue as a winged adult. 
During the nymphal life wings have been slowly developing, visible as 
short pads projecting from the dorsal margins of the meso- and meta-thorax, 
and appearing visibly larger after each moulting (Fig. 102). Respiration is 
accomplished by flat, leaf-like gills (Fig. 102) (these do not appear in some 
species until after one or two moultings), arranged segmentally along the 
sides of the abdomen. The mouth-parts are well developed for biting 
and chewing, with sharp-pointed jaws (mandibles). During its aquatic 
life at the bottom of stream or pond the May- 
fly has to undergo all the vicissitudes of an 
exposed and protracted life; it is eagerly sought 
after by larger, fierce, predaceous insects, 
stronger of jaw and swifter than itself; it is 
the prized food of many kinds of fishes, and it 
has to struggle with its own kind for food and 

At the end of the immature life the nymphs 
rise to the surface, and after floating there a 
short time suddenly split open the cuticle along 
the back and after hardly a second's pause 
expand the delicate wings and fly away. Some 
nymphs brought into the laboratory from a 
watering- trough at Stanford University emerged 
one after another from the aquarivim with 
amazing quickness. Almost all other insects 
require some little time after the final moulting 
for the gradual unfolding of the wings, and dry- 
ing and strengthening of the body-wall, before 
flight or other locomotion. Most of the May- 
fly species go through another moulting after 
acquiring wings, a phenomenon not known to occur in the case of any other 
insect. The stage between the first issuance from the water with expanded 
wings and the final moulting is called the subimago stage, and may last, 
in various species, from but a few minutes to twenty-four hours. Such 
is, in general, the life-history of the May-flies. As a matter of fact, the 
life-history of no single May-fly species has yet been followed completely 

Fig. 102. — Young (nymph) of 
May-fly, showing (g) tracheal 
gills. (After Jenkins and 
Kellogg; three times nat- 
ural size.) 


The May-flies and Stone-flies 

through. And here is an opportunity for some keen-eyed amateur ento- 
mologist to add needed facts to our knowledge of insect life. 

The breathing-organs of the nymph are of interest, as special adaptations 
to enable them to take up oxygen and give off carbon dioxide without com- 
ing to the surface, as do the water-beetles, water-bugs, mosquito-wrigglers, 
and many other familiar aquatic insects. Each plate-like gill (Fig. 102) 
is a flattened sac, with upper and lower membranous walls which run into 
each other all around the free margin. Inside this sac is an air-tube 

, (tracheal trunk) with numer- 
ous fine branches. By osmosis 
an interchange of gases takes 
place through the walls of the 
tracheae and of the sac — car- 
bonic dioxide passing out, and 
air from that held in solution 
in the water passing in. If a 
nymph held in a watch-glass 
of water be watched, at times 
all the gills will be seen rap- 
idly vibrating, thus setting 
up currents and bringing fresh 
aerated water to bathe the 

In the adult wihged stage 
(Fig. 103) the May-flies are 
extremely frail and delicate- 
bodied. The wings are fine 
and gauzy, consisting of 
the thinnest of membranes 
stretched over a perfect net- 
FiG. 103. — May-fly, from California. (Natural size.) ^ork of veins The fore 

wings are always markedly larger than the hind wings; in some species 
the latter are very small indeed, or even wanting altogether (Fig. 104). 
The body-wall is weakly chitinized, and collected specimens almost always 
shrivel and collapse badly in drying. The abdomen usually bears two 
or three long filaments on its tip; the head is provided with compound eyes 
and short awl-like antennte. The often-repeated statement in text-books 
that adult May-flies have no mouth nor mouth-parts is not literally true 
of all species, as weakly developed jaws and lips are present in some. But 
they are in such weak and atrophied condition that they can hardly be func- 
tional. It is probable, therefore, that no adult May-fly takes food. In 
the males of some species the compound eyes present a very interesting 

The May-flies and Stone-flies 


condition, being divided, each into two parts, by a narrow impressed line 
or by a broader space (Fig. 105). The two parts differ in the size of the 
facets of the ommatidia, i.e., eye-elements, and it has been ascertained (Zim- 
merman, 1897) tl^^t this difference in size of facets 
is accompanied by other and more important 
structural differences, which make it certain that 
the two parts of the eye have different powers of 
seeing. One part is especially adapted for seeing 
in the dark, or for detecting slight differences in 
intensity of light, but is ill-fitted for exact sight, 
while the other part is adapted for seeing in 
daylight, and for making a more exact picture of 
outhne. As the mating flights occur usually at 
twilight or in the evening, Zimmerman believes 
that this modification of the eyes of the males 
is to enable them to discover the females in the 
whirling shadow-dances. Chun has recorded a 
similar division and difference in the eye of 
certain ocean crustaceans and believes that the 

"dark eyes" are used for seeing in the dimly Fig. 104.— May-fly, Canis 
,. 1 , , , , ,, r 1-1 , /,,. 1 dimidiata, possessing only 

hghted water below the surface, while the light p - 

one pair of wings. 


eyes" are for special use at the brilliantly lighted 
surface. I have noted similar conditions in the eyes of both male and 
female net-winged midges (Blepharoceridae), small, two-winged flies of 
particularly interesting hfe (see p. 319). It is unusual to find such parallel 
adaptations in forms so unrelated. 

The May-flies show an anatomical condition of much interest to ento- 
mologists in the paired openings for 
the issuance of the eggs. Insects have 
their organs arranged in pairs, one on 
each side of the middle line of the 

f^^^T^ ///^ ' — ^"^^K ■ ^'^^^ body, as the legs, wings, mouth-parts, 
%^>^^^^'^^^^—^^^M^M antennae, eyes, spiracles, etc., or exact- 
^'^ ^"^W^ \y Qj^ ^]^g middle line, as the heart, 

alimentary canal, and ventral nerve- 
cord. That is, the typical insect body 
is bilaterally symmetrical, and the 
more apparent this symmetry is the sim- 
pler and more generalized the insect 
is believed to be. All other insects but the May-flies have the two egg- 
ducts, one from each egg-gland, fused inside the body, so as to form a short, 
single, common duct on the median line. But the May-flies have the ducts 

FiL.. 105. — Section through head of 
male May-fly, Potamanihus hriinneus, 
showing composition of compound 
eye and two sizes of eye-elements 
(ommatidia). (After Zimmer; greatly 

70 The May-flies and Stone-flies 

separate; that is, paired and bilateral for their whole course. This is taken 
to be an indication of the primitiveness and antiquity of the order. 

If the May-fiies are an ancient group of insects, and there is httle doubt 
of this, we have in them another example (we have previously noted one 
in the case of Campodea, see p. 60) of primitive insects of excessively 
frail and defenceless character persisting in the face of the strenuous struggle 
for existence and of the competition, in this struggle, of highly developed, 
specialized insect forms. Perhaps the solution of this problem in the case 
of the May-flies is to be found in their extreme prolificness and in the 
ephemeral character of their adult hves. It is only in the adult condition 
that May-flies are so ill-fitted to defend themselves; so they simply make no 
attempt to do so. They lay their eggs immediately on coming of age, and 
thus accomphsh the purpose of their adult stage. In their immature form 
they are not so handicapped in the struggle for existence, although they 
seem by no means in position to compete with some of their neighbors, like 
the nymphs of the stone-fly and dragon-fly. 

About 300 species of Ephemerida are known, of which 85 occur in 
North America. Their classification has been comparatively httle studied 
and is a difficult matter for beginners. The differences among the adults 
are so slight, and the preserved specimens are so uniformly misshapen 
and dried up, that most of us will have to be satisfied with knowing that 
we have in hand a May-fly, without being able to assign it to its genus. 
Keys to the North American tribes and genera of May-flies may be found 
by the student who may wish to attempt the generic determination of his 
specimens, in a paper by Banks in the Transactions of the American Ento- 
mological Society, v. 26, 1894, pp. 239-259. 

There are better defined differences among the nymphs than among 
the adults, but unfortunately the nymphs have been as yet too little studied 
for the making out of a comprehensive key to the genera. Needham and 
Betten give an analytical table of genera of Ephemerid nymphs as far as 
known in the Eastern United States, in Bulletin 47 of the New York State 
Museum, 1901. 

On the under side of the same stones in the brook "riffles" where 
the May-fly nymphs may be found, one can almost certainly find the very 
similar nymphs (Fig. 106) of the stone-flies, an order of insects called 
Plecoptera. More flattened and usually darker, or tiger-striped with black 
and white, the stone-fly nymphs live side by side with the young May-flies. 
But they are only to be certainly distinguished from them by careful exam- 
ination. The gills of the immature stone-flies usually consist of single short 
filaments or tufts of short filaments rising from the thoracic segments, one 
tuft just behind each leg (Fig. 106), and not flat plates attached to the sides 

The May-flies and Stone-flies 


of the abdomen as in the IVIay-fly nymphs. The feet of the stone-flies have 
two claws, while those of the young May-flies have but one. The stone-fly 
nymph has a pair of large compound eyes, as well as three small simple eyes, 
strong jaws for biting and chewing (perhaps for 
chewing heir nearest neighbors, the soft-bodied, 
smaller May-fly nymphs!), and two slender back- 
ward-projecting processes on the tip of the abdomen. 
The legs are usually fringed with hairs, which makes 
them good swimming as well as running organs. 
The nymphs can run swiftly, and quickly conceal 
themselves when disturbed. 

All stone-fly nymphs, as far as known, require 
well aerated water; they cannot live in stagnant 
pools or foul streams. Needham says that a large 
number of the smaller species are wholly destitute 
of gills absorbing the air directly through the skin. 
Nymphs brought in from a brook and placed in a Fig 106.— Young(nymph) 
; f .,, .,, , -11 ,-r- 1 01 stone-flv, from Cali- 

vessel of still water will be seen with claws arnxed, fornia. (Twice natural 

vigorously swinging the body up and down, trying size.) 
to get a breath under the difficult conditions into which they have been 
brought. The food-habits are not at all well known: some entomologists 
assert that small May-fly nymphs and other soft-bodied aquatic creatures 
are eaten, while others say that the food consists of decaying organic matter. 
Here is another opportunity for some exact observation 
by the interested amateur. On the other hand it is per- 
fectly certain that the nymphs themselves serve as food 
for fishes. 

The fully worked-out life-history of no stone-fly seems 
to have been recorded. The eggs, of which 5000 or 6000 
may be deposited by a single female, are probably dropped 
on the surface of the water, and sink to the bottom 
after being, however, weU distributed by the swift current. 
Sometimes the eggs are carried about for a while by the 
female, enclosed in a capsule attached to the abdomen. 
The young moult several times in their growth, but 
probably not nearly as many times as is common among 
May-flies. When ready for the final moulting, the nymph 

Fig. 107. — Exuvia 
of nymph of stone 

fly. (Natural size.) crawls out on a rock or on a tree-root or trunk on the 
bank, and splitting its cuticle along the back, issues as a winged adult. 
The cast exuviae (Fig. 107) are common objects along swift brooks. 

The adults (Fig. 108) vary much in size and color, the smallest being 
less than one-fifth of an inch long, while the largest reach a length of two 

72 The May-flies and Stone-flies 

inches. Some are pale green, some grayish, others brownish to black. 
There are four rather large membranous, many-veined wings without pattern, 
the hind wings being larger than the front ones. When at rest, the fore 
wings He flat on the back, covering the much-folded hind wings. The mouth- 
parts are present and are fitted for biting, although the food-habits are not 
known. It is asserted that some species take no food. The antennae are 
long and slender. The abdomen usually bears a pair of long, many-seg- 
mented, terminal filaments. The body is rather broad and flattened, and 
there is no constriction between the thorax and abdomen. On the ventral 
aspect of each thoracic segment there is a pair of small openings whose func- 

FiG. io8. — A stone-fly, Perla sp., common about brooks in California. (After Jenkins 
and Kellogg; twice natural size.) 

tion is unknown. The adults of certain species retain, although in shriveled 
and probably functionless condition, the filamentous gills. This fact is of 
importance in connection with the question as to whether insects are 
descended from aquatic or terrestrial ancestors. Those who believe in 
the aquatic ancestry have found a simple origin for the spiracles (breathing- 
pores) by imagining them to be the openings left when the gills, used in 
aquatic life, were lost. But the adult stone-flies which retain their gills 
also have wholly independent spiracles. 

About ICO species of stone-flies are known in North America. The 
adults are to be found flying over or near streams, though sometimes 

The May-flies and Stone-flies 


straying far away. They rest on trees and bushes along the banks. The 
green ones usually keep to the green foliage, while the dark ones perch on 
the trunk and branches. The various species are included in ten genera, 
which may be determined by the following table: 


The following technical terms not heretofore defined are used in this key: cerci, 
slender processes projecting from the tip of the abdomen; radial sector, cubital vein, 
and other names of veins in the wings may be understood by reference to Fig. 109. 


Fig. 109. — Diagram of venation of wing of a stone-fly; /, costal vein; 2, subcostal vein; 
3, radial vein; 4, medial vein; 5, first anal vein; 6, radial sector, P, pterostigma; 
A, arculus: Op a^, a^, apical cells. Between the medial and first anal vein is the 
cubital vein, not numbered. Cell M is the cell behind the medial vein; cell Sc is the 
cell behind the subcostal vein. 

A. With two long, many-jointed cerci. 

B. Radial sector not reduced, i.e., with four or more branches. 

C. Wings strengthened throughout by many cross-veins, there being many 
cross-veins between the branches of the media, between the accessory 
cubital veins, and in the anal areas of both pairs of wings. .Pteronarcys. 
CC. Wings with few or no cross-veins between the branches of the media, 
between the branches of the cubital veins, and in the anal area. 
D. Radial area of the fore wings with an irregular network of veins- 


DD. Radial area of the fore wing with no cross-veins except the radial 
cross-veins, or with a few regular cross-veins. .. .Perla (in part). 
BB. Radial sector reduced, i.e., with less than four branches. 
C. Hind wings much broader than the fore wings. 

D. With several cross-veins in cell M of the fore wings. 

E. Cell Sc of the fore wings with at least three cross-veins. 

F. With three ocelli Perla (in part). 

FF. With only two ocelli Pseudoperla. 

EE. Cell Sc of the fore wings with only one or two cross-veins. 

Small species of a green or yellow color Chloroperla. 

DD, With only one cross-vein in cell AI of the fore wings between the 

arculus and the medio-cubital cross-vein Capnia. 

CC. Hind wings of the same width as the fore wings; the anal area of the 

hind wings not expanded Isopteryx. 

AA. With the cerci rudimentary or wanting. 

B. Second segment of the tarsi equal in length to the others; rudimentary cerci 
present T.eniopteryx. 

74 The May-flies and Stone-flies 

BB. Second segment of the tarsi small, shorter than the others, cerci absent. 

C. Veins radiating from the ends of the radial cross-vein forming an X. 

CC. Veins radiating from the ends of the radial cross-vein not forming an X. 


The genus Perla (Fig. io8) includes more species than any other. The 
species of Pteronarcys retain gills in the adult condition. The species of 
Chloroperla are small, delicate, and pale green. Leuctra includes the slender- 
est of the stone-flies; they are small and brownish. Comstock says that 
there are several species of stone-flies that appear on the snow on warm 
days in late winter. They become more numerous in early spring, and 
often find their way into houses. The most common one in Central New 
York is the small snow-fly, Capnia pygnma, which is grayish black. The 
female is 9 mm. (about f in.) long, with an expanse of wings of 16 mm. 
(about I in.), while the male is but 4§ mm. (about \ in.) long, and has 
short wings which extend but two-thirds the length of the abdomen. 


SEL-FLIES (Order Odonata) 

HEN it is high noon on the mill-pond, — 
when leaves droop, and sun glares upon the 
water, and the air is hot and still, when 
other creatures seek the shade, and even 
the swallows that skim the air morning 
and evening are resting, — then those other 
swallows of the insect world, the dragon- 
flies, are all abroad. . . . One may stand 
by the side of a small pond, and follow 
for hours with his eye the evolutions of one of the large dragon-flies skim- 

■l^-^^-lAlel l-n^ar 

ming over the surface in zigzag Hnes or sweeping curves, stopping still 
in midair, and starting again, seeming never to rest, nor even to tire. Poised 



Dragon-flies and Damsel^flies 

in the air, with the sunUght dancing on its trembhng wings, it is indeed a 

beautiful sight. 

"'Dragon-flies? Folks call 'em devil's-darnin'-needles in our parts, 

and they say they will sew up your ears.' Yes; and in some localities they 

are called 'snake-doctors,' and are said 
to bring dead snakes to life; and other 
meaningless names are given them, such 
as 'snake-feeders,' 'horse-stingers,' 'mule- 
killers,' etc.; but in spite of all these 
silly names and the silly superstitions 
they represent, dragon-flies are entirely 
harmless to man — are indeed to be 
counted as friends, for they destroy vast 
numbers of mosquitoes and gnats and 
pestiferous little flies. To such creatures 
they must seem real dragons of the air. 
While one is standing by the pond let 
him follow awhile the actions of a dragon- 
fly that is making short dashes in different 
Fig. 1 10.— a dragon-fly (from life), directions close to the bank. Let him 

fix his eye on a little fly hovering in the air, and note that after the 

dragon-fly has made a dart toward it, it is gone. Let him repeat the 

observation as the dragon-fly goes darting 

hither and thither. It will be hard to see 

the flies captured, so quickly it is done, 

but one can see that ' the place that once 

knew them knows them no more.' And 

the usefulness of the dragon-fly in taking 

off such water-haunting pests will be 


Thus entertainingly and truthfully writes 

Professor Needham of the strong-winged, 

brilliantly colored, graceful insects of our 

present chapter. If one could see through 

muddy water and would fix his gaze on 

the weed-choked slimy depths of the pond, Fig. m 

he would see the dragon-flies in another 

stage of their life, under very different 

conditions of existence, and in very different guise. Crawling awkwardly 

about over and through the decaying weeds and leaves and mud of the 

bottom or lying in ambush, half concealed by coverings of slime, 

would be seen certain strange big-headed, thick-bodied, dirty gray-green, 

The young (nymph) of 
a dragon-fly. (From Jenkins and 
Kellogg; twice natural size.) 

Dragon-flies and Damsel-flies 'j'j 

wingless creatures from half an inch to two inches long. Occasionally 
one of these creatures suddenly darts forward by spurting water from 
the hinder tip of its body; occasionally one quickly thrusts out from 
its head a vicious pincer-like organ which is more slowly withdrawn, or 
rather folded up, with an unfortunate tiny water-animal squirming in the 
toothed pincers. Still dragons, though now dragons of the deep instead of 
flying dragons, these are our insects in their immature or larval life. Their 

Fig. 112. — Young (nymph) dragon-fly, showing lower lip folded and extended. (From 
Jenkins and Kellogg; twice natural size.) 

prey, consistmg of water-bugs, May-fly larvae, small crustaceans, mol- 
lusks, and any of the numerous aquatic insect larvae, including other 
young dragon-flies, is probably always caught alive. Not by active 
pursuit, as in the air above, but by lying in wait in the murky depths 
of the pond until the unsuspecting insect comes within reach of the 
extensible lower lip with its pair of broad spiny, jaw-like flaps at the 
clutching tip. The fierce face of the young dragon, with its great 
mouth and sharp jaws, is all concealed by this lip when folded up, 
and there is little in the appearance of the dirty, sprawling, smooth- 
faced creature to betray its dragon-like character. But appearances in 
the insect world may be as deceptive as in our own, and too late the 
careless water-bug out on a foraging swim for lesser prey finds himself in 
range of a masked battery and becomes the preyer preyed upon. 

About three hundred different species of dragon- and damsel-flies 
(damsel-flies are the smaller, slender-bodied, narrow-winged kinds, see Fig. 
113) are known in North America, about two thousand having been found 
in all the world. In any single locality where conditions are at aU favor- 
able to dragon-fly life, that is, where there are live streams and ponds, from 
a score to two or three times as many different dragon-flies can be found. 
One hundred species occur in Ohio, and one hundred and twenty in New 
York, states offering specially favorable natural conditions for them, while 
only about fifty species have been found in California, a much larger but 
more arid region. The young of no dragon-fly species is known to live in 
salt water, although nymphs have been found in brackish water and in 


Dragon-flies and Damsel-flies 

streams impregnated with sulphur from sulphur springs. Nor do dragon- 
flies like cold weather. Although a few species are found in the far North 
(recorded at 70° N. in Norway, 65° N. in Alaska, and 63° N. in Siberia) 
and a few at high cold altitudes (as high as 10,000 feet) on mountain flanks, 
the great majority of them need considerable temperature for growth and 
development and even for activity during adult life. Calvert says that but 
one species is known which regularly passes the winter in adult stage, and 

that most dragon-flies live as adults from 
but twenty-five to forty-five days, and 
these in the summer. In California, where 
the winter temperature at sea-level only 
occasionally falls to 32° F., adult dragon- 
flies can be found in most of the months 
of the year. 

The adult dragon-flies are to be seen 
pursuing their prey, like hawks, with 
swift darting flights over ponds, along 
streams, and even scattered widely inland 
over fields and in woods. A few kinds 
have a liking for the vicinity of houses. 
Needham, a careful student of these 
insects, has found that the hunting region 
above and along the shores of a pond may 
be imaginarily divided into zones one 
above the other, each zone characterized 
by the presence of a few particular 
dragon-fly species. "So, in fact," he writes, "we find the smaller damsel- 
flies flying over the water in a straight course an inch or less above the 
surface, and rarely venturing higher; the larger damsel-flies a little higher; 
the amber wings at an average of about six inches; the larger skimmers 
a foot or more from the surface, and upland skimmers and darters still 
higher. One has only to stand a little while by some small area of water 
where all these are flying to see that each keeps rather closely to his proper 
altitude. Why do damsel-flies keep so close to water? The reason is 
not far to seek. Dragon-flies eat one another — the strong destroy the 
weak. If to venture up into the altitude of the larger species means to run 
the risk of being eaten, we can readily see why the damsel-flies should 
stay down below. The hawk may roam the air at will, but sparrows must 
keep to the bushes." 

We think of dragon-flies, as of albatrosses and Mother Carey's chickens, 
as being always on the wing. They catch their prey while flying, eat it 
while flying, mate while flying, and some of them deposit their eggs while 

Fig. 113. — Damsel-flies 
winged dragon -flies), 
size; from life.) 


Dragon-flies and Damsel-flies 79 

on the wing. But of course all dragon-flies rest sometimes, and some of 
them, especially the damsel-flies, are at rest most of the time, cHnging to 
stems or leaves by the water's edge. The larger kinds may be found 
occasionally perched on the tips of tall swaying reeds, or on a stump or 
projecting dead limb. From these coigns of vantage they swoop like 
a hawk on any rash midge that ventures awing in the neighborhood. 
Cold or cloudy weather, or a strong wind, will drive most dragon-flies to 

The Odonata are unexcelled among insects for swiftness, straightness, 
and quick angular changes in direction of flight. The successful main- 
tenance of their predatory life depends upon this finely developed flight 
function together with certain structural and functional body conditions 
which might be said to be accessory or auxiliary to it. And this may be 
an appropriate place to describe briefly a few of their sahent structural 

All dragon-flies have four well-developed wings, and all show such a 
similar general bodily make-up and appearance, that from an acquaintance- 
ship with two or three familiar species any member of the order can be 
recognized as really belonging to the group. The body in all is long, smooth, 
and subcylindrical or gently tapering. This clean, slender body offers 
little resistance to the air in flight, and serves as an effective steering-oar. 
The wings are long and comparatively narrow, fore and hind wings being 
much alike, almost exactly ahke indeed in the damsel-flies. The venation 
is of the general type known as net-veining (Fig. 114b), the few strong longi- 
tudinal veins being connected by many short cross-veins. The fore wings 
are greatly strengthened along their costal (front) margin by having the 
first longitudinal (subcostal) vein behind the margin placed at the bottom 
of a groove, and the cross-veins in that groove so enlarged vertically as 
to take on the character of flat, plate-like braces or buttresses. As, in 
the figure-of-eight movement of the wing in flight, the front margin first 
meets the resistance of the air, it is necessary that swiftly and strongly beat- 
ing wings should be especially strengthened along this edge, and this is just 
what the peculiar folding and bracing of the costal region of the dragon-fly's 
fore wing accomplishes. 

The head is unusually large and is more than two-thirds composed of 
the pair of great compound eyes. More than 30,000 facets have been 
counted in the cornea of certain dragon-fly species, and this means that each 
eye is made up of more than 30,000 distinct eye-elements or ommatidia, 
each capable of seeing a small part or point of any object in range of vision. 
Thus an image of a near-by object is made in fine mosaic, and the finer the 
mosaic the more definite and precise is the vision by means of compound 
eyes. These great eyes, too, have facets directed up and down and sidewise 

8o Dragon-flies and Damsel-flies 

as well as forward, and by a special sort of articulation of the head on the 
thorax it can be rotated readily through i8o°, so that the principal part of 
each eye can be directed sidewise or even straight down. For accurate 
flight and successful pursuit of flying prey the dragon-fly has full need of 
good eyes. It is to be noted, too, that the eyes are relatively largest in those 
particular dragon-fly kinds which have the most powerful flight. On the 
head, also, are three simple eyes (ocelli), the pair of very small awl-like 
antennae, and the great mouth. The mouth is overhung as by a curtain 
by the large flap-like upper lip (labrum). The jaws (mandibles) are strong 
and toothed, and obviously well adapted for tearing and crushing the cap- 
tured prey. 

When the prey is come up with, however, it is caught not by the mouth 
but by the "leg-basket." The thorax is so modified, and the insertion of 
the legs such, that all the legs are brought close together and far forward, 
so that they can be clasped together like six slender, spiny grasping arms 
just below the head. Although the catching and eating is all done in the 
air and very quickly, observers have been able to see that the prey is caught 
in this "leg-basket" and then held in the fore legs while being bitten and 
devoured. These slender legs are used only very slightly for locomotion, 
but they serve well for the light unstable perching which is characteristic 
of the dragon-flies.. 

The internal anatomy is specially characterized, as might well be 
imagined, by a finely developed system of thoracic muscles for the rapid 
and powerful motion of the wings and the delicate and accurate move- 
ments of the legs. The respiratory system is also unusually well developed, 
such active insects needing a large quantity of oxygen, and generating a 
large amount of carbon dioxide. The respiratory movements, according 
to Calvert, consist in an alternate expansion (inspiration through the ten 
pairs of breathing-holes, or spiracles, arranged segmentally on thorax and 
abdomen) and contraction (expiration) of the abdomen. The rate of 
movement varies greatly at difl'erent times owing to unknown causes, but 
is always quickened by exercise, increased temperature, or mechanical irri- 
tation. In different dragon-flies the inspirations have been noted to be 
from 73 to ii8 a minute. 

The dragon-flies are famous for their beautiful metallic colors. As they 
dart through the air one gets glimpses of iridescent blue and green and cop- 
per, of tawny red and violet and purple reflections that are most fascinating 
and tantalizing. Seen close at hand in the collections, however, they are 
mostly dull-colored and, except for their "pictured" wings and the sym- 
metry and trim outline of their body, rather unattractive "specimens." But 
a freshly caught dragon-fly shows the real glory of the coloring: delicate 
changing shades of green and violet and copper quiver in the great eyes; 

Dragon-flies and Damsel-flies 8 1 

the thorax is transkicent green or blue, and the long symmetrical body is 
warm red or deep blue or purple or green. It is often covered with a soft 
whitish "bloom," that tones down the brilliant metallic iridescence. But 
as the body dries, the colors fade. They are due not so much to pigment 
as to the interference in reflection of the various color-rays, this interference 
being caused by the structure of the body-wall. Just as soap-bubbles or 
weathered plates of glass or mica produce brilliant colors by interference 
effects, so does the semi-transparent laminate outer body-wall of the 
dragon-fly produce its fleeting color glories. While the wings of many 
kinds are clear, unmarked by blotches or line, the wings of others bear a 
definite "picture" or pattern, usually light or dark brown or even blackish, 
reddish, thin yellow, or whitish. These wing-patterns make the determination 
of many of the dragon-fly species a very simple matter. 

When the dragon-flies go winging about over ponds and streams they 
are engaged in one of three things: in eating, in mating, or in egg-laying. 
The prey of the dragon-fly may be almost any flying insect smaller than 
itself, although midges, mosquitoes, and larger flies constitute the majority 
of the victims. Howard says that the voracity of a dragon-fly may easily 
be tested by capturing one, holding it by its wings folded together over its 
back, and then feeding it on Hve house-flies. Beutenmiiller found that 
one of the large ones would eat forty house-flies inside of two hours. Howard 
says that a dragon-fly will eat its own body when offered to it (query, to 
its head?) and that a collected dragon-fly, if insufficiently chloroformed and 
pinned, will when it revives cease all efforts to escape if fed with house-flies, 
the satisfying of its appetite making it apparently oblivious to the discom- 
fort or possible pain of a big pin through its thorax. That dragon-flies 
are sometimes cannibalistic has been repeatedly confirmed by observation. 
The nymphs have been seen to devour nymphs of their own and other 
species; the nymphs of a European form have been observed to come out of 
water at night and attack and devour newly transformed imagoes of the 
same species, while several instances are recorded of the capture and devouring 
of an imago of one species by an imago of another. 

The good that is done by dragon-flies through their insatiable appetite 
for mosquitoes is very great. Now that we recognize in mosquitoes not 
only irritating tormentors and destroyers of our peace of mind, but alarm- 
ingly dangerous disseminators of serious diseases (malaria, yellow fever, 
filariasis), any enemy of them must be called a friend of ours. A prize was 
once offered for the best suggestions looking toward practicable means of 
artificially utilizing dragon-flies for the destruction of mosquitoes and house- 
flies, but no very efficient improvement on the dragon-fly's natural tastes 
and practices were brought out by this essay competition. 

In Honolulu, the principal city of our mid-Pacific territory, the mosqui- 

82 Dragon-flies and Damsel-flies 

toes are so abundant that no one neglects to enclose his bed carefully each 
night in mosquito-netting, and all bedrooms are equipped with an ingenious 
canopy which can be folded closely in the daytime and readily spread over 
the bed at night. The continuous and abundant presence of mosquitoes 
is such a matter of fact that it has dictated certain particular habits of life 
to the inhabitants of Honolulu. But in the daytime one is singularly free 
from mosquito attack. Coincidentally with this one notes the surprising 
abundance and strangely . domestic habits of great dragon-flies. I have 
watched dozens of dragon-flies hawking about a hotel lanai (porch) in the 
heart of the town. No pond or stream is nearer than the city's outskirts. 
Dragon-flies are in the main streets, in all the gardens, and they are chiefly 
engaged in the laudable business of hunting the hordes of "day" mosquitoes 
to their death. The most conspicuous features of insect life in Hawaii are 
the hosts of dragon-flies by day and the hordes of mosquitoes by night. As 
the dragon-flies unfortunately are not night flyers (although some forms 
keep up the hunting until it is really dark), it is by night that one realizes 
what a plague the mosquito is in the islands. Were it not for the dragon- 
flies, life in the islands would be nearly intolerable. The rice-swamps and 
taro-marshes and the heavily irrigated banana and sugar plantations offer 
most favorable breeding-grounds for the mosquitoes, but also fortunately 
for the dragon-flies as well. The mosquitoes of Hawaii are not indigenous; 
they were introduced with white civilization. It is told, and is not improb- 
able, that the skipper of a trading schooner in early days, to revenge himself 
for some slight put on him by the natives, purposely put ashore a cask of 
water swarming with mosquito wrigglers. It needed no more than that 
to colonize this fascinating tropic land with the mosquito plague. How 
the saving dragon-flies came is not yet come to be tradition; indeed, few 
Hawaiians understand how important a part the dragon-fly plays in their 
life. They do appreciate the mosquito. 

In the Samoan Islands, too, where we have another tropical colony, 
the mosquitoes are a great plague. Here the matter is made more serious. 
The Samoan mosquitoes are carriers and disseminators of a dreadful disease 
known as elephantiasis from the enormous enlargement of the legs and 
arms of sufferers from it. This disease is the great scourge of these islands, 
more than 30% (from my own observation; 40% and 50% are estimates 
given by other observers) of the natives having it. (For an account of 
the role of mosquitoes in the dissemination of malaria, yellow fever, and 
elephantiasis, see Chapter XVIII of this book.) The dragon-flies are, in 
Samoa as in Hawaii, conspicuous by their abundance and variety, and they 
do much to keep in check the quickly breeding mosquitoes. 

Watching the flying dragon-flies over a pond, you may occasionally 
see one poising just over the surface of the water, and striking it with the 

Dragon-flies and Damsel-flies 


tip of the abdomen; or another kind may be seen to swoop swiftly down to 
the surface occasionally in its back-and-forth flight, and to dip the tip of 

Fig. 114a. 

Fig. 1 146. 
Stages in the development of the giant dragon-fly, Anax Junius, a, youngest stage; b, 
c, and d, older stages, showing gradual development of the wings. (Young stage, 
slightly enlarged after Needham; adult three-fourths natural size.) 

the body for a moment into the water. These are females engaged in laying 
their eggs. The eggs issue in small masses, usually held together by a gelat- 
inous substance. From several hundred to several thousand eggs are laid by 


Dragon-flies and Damsel-flies 

each female. Needham counted 110,000 eggs in a single egg-mass of Libellula. 
Sometimes the eggs may be laid on wet mud or attached to moist water- or 
shore-plants. The damsel-flies and a few of the dragon-flies insert the eggs 
in the stems of dead or living water-plants below the surface of the water. 
To do this they have to cling to the stem, with the abdomen or sometimes 
the whole body under water, and cut slits in it with the sharp ovipositor. 
The eggs are sometimes laid on submerged timbers and moss- or alga-covered 
stones. Kellicott observed females of A rgia putrida (a damsel-fly abundant 
along Lake Erie) to remain wholly under water for from five to fifty-five 
minutes at a time. These females were accompanied by males which also 
stayed under for similar lengths of time. 

The eggs hatch after various periods, depending on the species of dragon- 
fly and on the time of year of oviposition. In midsummer Needham found 
the eggs of some species to hatch in from six to 
ten days, while others laid in autumn did not hatch 
until the following spring. In the same lot of eggs 
the period of incubation may vary even in midsum- 
mer from a week to more than a month. 

From the eggs come tiny, spider-like nymphs 
with long, slender legs, thin body, and no sign of 
wings. Even in the largest dragon-fly species the 
just-hatched young are only about one-twelfth of 
an inch long, while the nymphs of the common 
Libellulas are only one-twenty-fifth of an inch long 
at hatching. They begin their predatory life, con- 
fining their attention at first to the smaller aquatic 
creatures, but with increasing size and strength 
and confidence being ready to attack almost any of 
the under-water dwellers. Even fish are seized by 
the larger nymphs, Needham having seen the 
nymphs of one species seize and devour young 
brook-trout as long as themselves. 

The young of different species differ consider- 
ably in size, shape, and duration of their nymphal 
existence. The nymphs of some species require 
more than a year to develop into adults, while those of some others are ready 
to transform in a few months, not a few dragon-fly species having two gener- 
ations a year. The one-year life cycle, however, is usual among the more 
familiar dragon-flies, the eggs laid during midsummer hatching in late sum- 
mer, the nymphs hibernating and being ready to emerge the following sum- 
mer. Needham thinks that the damsel-flies have a number of broods in 
a season, the processes of transformation and oviposition beginning as soon 

Fig. 115. — The young 
(nymph) of a damsel- 
fly (narrow-winged dra- 
gon-fly), Lestes sp. The 
three leaf -like processes 
at the tip of the abdo- 
men are gills (Twice 
natural size.) 

Dragon-flies and Damsel-flies 85 

as the weather permits, and continuing industriously to the close of the 

The nymphs cast the skin repeatedly during their growth and develop- 
ment, although the exact number of moultings is not known for any species. 
After two or three moults the wing-pads appear and with each successive 
moult increase in size. Immediately after moulting the nymphs are Ught 
greenish or gray, and their characteristic color pattern is distinct, but they 
gradually darken, the pattern becoming more and more obscure until by 
the t'me for another moulting the body is uniformly dark and dingy. The 
nymphs (Fig. 115) of the damsel-flies are elongate and slender, and have 
three long conspicuous gill-plates at the tip of the abdomen, which they 
can also use as sculls for swimming. The dragon-fly nymphs are robust- 
bodied, some of them indeed having the abdomen nearly as wide as long 
and much flattened. All the nymphs are provided with the long grasping 
lower lip, which can be folded mask-like over the face when not engaged 
in seizing prey. The mandibles are strong and sharp and the whole mouth 
is well fitted for its deplorable but necessary business. 

The true dragon-fly nymphs do not have plate-like gills, like those of the 
damsel-flies, nor any other external kind, but have the posterior third of 
the intestine lined with so-called internal gills. These internal or rectal 
gifls are in six longitudinal bands, each consisting of two thin rows of small 
plates or tufts of short slender papillae. Water is taken into the intestine 
through its posterior opening and, after bathing the gills, giving up its dis- 
solved oxygen, and taking up carbon dioxide, it is ejected through the same 
opening. When this water is ejected violently it serves to propel the nymph 
forward. It is also apparently occasionally used for defence. 

Just as the adult flying dragon-flies keep to certain regions above or 
in the neighborhood of the pond, so Needham has found the nymphs to 
have various preferred lurking-places in the pond. The damsel-fly nymphs 
and a few of the more active dragon-fly nymphs clamber among submerged 
vegetation or inhabit driftwood and submerged roots or brush. The heavier 
sprawling Libellulid nymphs usually crawl over the bottom or climb over 
fallen rubbish, while certain other Libelluhds and some similar forms occupy 
the mud or sand of the bottom. The nymphs of one of these latter kinds 
is described as each scratching a hole for itself and descending into it like 
a chicken into a dust-bath, kicking the sand over its back and burrowing 
until all but hidden, only the tops of its eyes, the tips of its treacherous labium, 
and the respiratory aperture at the end of the abdomen reaching the surface. 

After the few weeks or month or year which the nymph requires for its full 
growth and development it is ready to transform. If in early summer, when 
the dragon-flies are beginning to appear, one will go out to the dragon-fly 
pond a little after daylight, he will see this transforming or issuance of the 


Dragon-flies and Damsel-flies 

winged imagoes busily going on. The nymphs crawl out of the water, and 
up on stones or projecting sticks, or on bridge-piles or the sides of boats, 
or on the stems of weeds growing by the water's edge. Here they cling quietly, 

awaiting the moment when the chi- 
tinous body-wall shall split lengthwise 
along the back of the thorax, and the 
made-over body inside with its damp, 
compressed wings, its delicate trans- 
parent skin, and changed mouth-parts 
and legs shall slowly work its way out 
of the old nymphal coat. The nymphs 
of some dragon-flies and damsel-flies 
crawl out among the weeds and grass 
of the shore for some distance before 
choosing a resting-place, and none of 
these will be very readily seen. But careful searching in a place from which 
winged individuals are occasionally arising will soon reveal the transforming 
in all of its stages (Fig. ii6). It takes some time for the emergence of the 
damp, soft imago from the nymphal skin, and some further time for the 
slow expanding and drying of the wings, and the hardening of the body- 
wall so that the muscles can safely pull against it. When all this has come 
about the imago can fly away. But even yet the colors are not fully acquired 

Fig. ii6. — The issuance of an adult white 
tail, Plathemis trimaculata. (After Need- 
ham; natural size.) 

Fig. 117. — Adult and last exuvia of the whitetail, Plathemis trimaculata. 
(Natural size.) 

and fixed, and these fresh imagoes have an unmistakably new and shiny 
appearance. They are called teneral specimens. Usually the emergence 
of nymphs from the pond and the subsequent transforming cease by the 
middle of the forenoon, and after that one can find only the frail, drying 

Dragon-flies and Damsel-flies 


Adult and last exuvia of the damsel- 
fly, Lestes uncata. (Natural size.) 

cast nymphal skins or exuvicX, clinging here and there to stones and plant- 
stems. Attached to these exuvice there may be often noted two or three short, 
white, thread-like processes. These 
are the dry chitinous inner linings 
of the main tracheal trunks of the 
dragon-fly which were moulted with 
the outer body-wall. As the main 
tracheal tubes are really invagina- 
tions of the outer skin, it is obvious 
that the inner lining of the trachea 
is continuous with the outer coat 
(chitinized cuticle) of the body-wall 
and so is naturally cast off with it. 
Although the habits of the adult 
dragon-flies must be studied out of 
doors, the nymphs can be brought 
indoors and kept alive so that their 
walking and swimming and hiding Fig. 118 
and capturing of prey, and often 
their transformation into winged imagoes, can be readily observed. In 
their natural habitat some of these observations are nearly impossible, 

and for school-room or private-study aquaria 
hardly any other animals can be found of 
more interest to the observer, whether child or 
grown-up, than the dragon-fly nymphs. 

Professor Needham, who has done more 
and better work in the study of the immature 
life of dragon-flies than anybody else, gives 
the following directions for collecting and 
rearing the nymphs: 

"If one wishes to collect the nymphs which 
lie sprawling amid fallen trash, a garden-rake 
with which to draw the trash aside, fingers not 
too dainty to pick them up when they make 
themselves conspicuous by their active efforts 
to get back into the water, and a pail of 
water in which to carry them home, are all 
-A home-made water- the apparatus required, 
collecting dragon-fly "A rake will bring ashore those other 
(After Needham.) ^y^^^is which burrow shallowly under the 
sediment that lies on the bottom, and also a few of those that cling to vegeta- 
tion near the surface; but for getting these latter a net is better. Fig. 119 

Fig. iiq.- 
net for 


Dragon-flies and Damsel-flies 

shows the construction of a good water-net that can be made at home out 
of a piece of grass-cloth, two sizes of wire, and a stick. 

"The best places to search for dragon-fly nymphs in general are the 
reedy borders of ponds and the places where trash falls in the eddies of 
creeks. The smaller the body of water, if permanent, the more likely it 
is to yield good collecting. The nymphs may be kept in any reasonably 
clean vessel that will hold water. Some clean sand should be placed in 
the bottom, especially for burrowers, and water-plants for damsel-fly nymphs 
to rest on. They may be fed occasionally upon such small insects (smaller 
than themselves) as a water-net or a sieve will catch in any pond. Their 
habits can be studied at leisure in a dish of water on one's desk or table. 

"The best season for collecting them is spring and early summer. April 
and May are the best months of the year, because at this time most nymphs 

are nearly grown, and, if taken then, 
will need to be kept but a short time 
before transforming into adults. And 
this transformation every one should 
see; it will be worth a week's work at 
the desk; and as it can be appreciated 
only by being seen, some simple direc- 
tions are here given for bringing the 
Fig. 1 20.— a simple aquarium for rear- ^ymphs to maturity. Place them in a 
ing dragon-fly nymphs. (After Need- •' ^ •' 

ham.) wooden pafl or tub (Fig. 120). if 

the sides are so smooth that they cannot crawl up to transform, put some 
sticks in the water for them to crawl out on. Tie mosquito-netting tightly over 
the top, or, better, make a screen cover; leave three or four inches of air 
between the water and the netting; feed at least once a week, set them where 
the sun will reach them; and after the advent of warm spring weather look 
in on them early every morning to see what is going on." 

Elsewhere Professor Needham says that nymphs may be fed bits of 
fresh meat in lieu of live insects. If meat is fed, it must be kept in motion 
before them, as they will refuse anything that does not seem alive. Some 
nymphs will take earthworms. Care must be taken to keep cannibalistic 
kinds apart from others. When the nymphs transform the freshly issued 
imagoes should be transferred each with its cast skin (exuvia) to dry boxes 
for a short time, till their body-wall and wings gain firmness and the colors 
are matured. The imago and its exuvia should always be kept together. 

Specimens of the adults for the cabinet should have the wings spread 
like butterflies and moths (for directions for spreading see the Appendix). 
The slender and brittle dried abdomen breaks off very easily, and a bristle 
or fine non-corrosive wire should therefore be passed lengthwise through 
the body as far as the lip of the abdomen. A couple ot insect-pins, inserted 

Dragon-flies and Damsel-flies 89 

lengthwise one at each end of the body, are used by some. Specimens 
intended for exchange should not be pinned up, but "papered," i.e., put 
with folded wings into an enclosing little triangular paper envelope made 
by folding an oblong paper sheet once diagonally and then folding over 
slightly the two margins. 

i %v / /■ / ■■■ A. P 

Fig. 121. — Diagram of venation of wing of dragon-fly. a, antecubitals; b, postcubitals; 
N, nodus; P, pterostigma; A, arculus; t, triangle. (After Banks.) 


Key to Suborders (Imagoes). 

Front and hind wings nearly similar in outline, and held vertically over the back 
when at rest; head wide and with eyes projecting and constricted at base. 

(Damsel-flies.) Suborder Zygoptera. 

Front and hind wings dissimilar, hind wings usually being much wider at base, and 

both pairs held horizontally outstretched when at rest; eyes not projecting 

and constricted at base (Dragon-flies.) Suborder Anisoptera. 

Key to Suborders (Nymphs). 
Posterior tip of abdomen bearing three, usually long, leaf-like tracheal gills. 

(Damsel-flies.) Suborder Zygopter.a. 
Posterior tip of abdomen with five, converging, short, spine-like appendages. 

(Dragon-flies.) Suborder Anisoptera. 

Key to Families (Imagoes). 

Wings with not less than five antecubital cross-veins (Fig. 121). 

Family Calopterygid^. 
Wings with not more than three, usually two, antecubitals (Fig. 121). 

Family Agrionid,e. 
Key to Families (Nymphs). 

Basal segment of the antennae extremely elongate Family Calopterygid.e. 

Basal segment of the antennae short, subrotund Family Agrionid^. 

The family Calopterygidae includes but two genera, Calopteryx, in which 
the basilar space of the wings is open and the wings themselves are rather 
broad near the tip, and Hetaerina, in which the basilar space is net-veined 
and the wings narrow. 

Calopteryx maculata (Fig. 122), the most familiar representative in the 
Eastern States of the first genus, has velvety black spoon-shaped wings, 


Dragon-flies and Damsel-flies 

Fig. 122. 

-The black wing, Calopteryx 
macula ta. 

(brownish in freshly moulted, or teneral specimens), and a long, slender body, 
of striking metallic blue or green. The females can be distinguished from 
the males by their possession of a milk-white pterostigma (Fig. 121). These 
beautiful "black wings" are found 'n gentle fluttering flight, usually along 
small streams in woods or meadows. The female lays her eggs "among 

the rubbish and mud along the 
borders of ditches," and the 
nymphs found in the ditches 
and streamlets have the middle 
one of the three caudal gills fiat 
and shorter than the other two. 
Kellicott has seen the males of 
this species fight fiercely with 
each other. "Two will fly about 
each other, evidently with con- 
suming rage, when one finally 
appears to have secured a posi- 
tion of advantage and darts at 
his enemy, attempting, often suc- 
cessfully, to tear and damage 
his wings." 

The best known representative of the other genus is a perfect master- 
piece of insect beauty and grace. Entomologists know it as Hetcerina 
americana (Fig. 123); I suggest that we call it the "ruby-spot," although 
only the males bear the gem. The head and thorax of the males are 
coppery red, the abdomen me- 
tallic green to coppery, and the 
basal fourth of each of the long, 
slender, and otherwise clear wings 
is bright blood-red. In the females 
the whole body is metallic green, 
with the basal third of the wings 
pale yellowish brown. These dam- 
sel-fiy beauties are shy and retiring, 
rarely venturing more than a few 
feet away from the willow-overhung 
bank of their favorite swift-running 
stream. Sometimes hundreds of 
them come together and chng in 
graceful festoons to the drooping willow branches. Then they look like 
strings of rubies, or of warm red flowers or seeds. 

The family Agrionidae includes the host of slender-bodied, narrow- and 

Fig. 123. — The ruby-spot, Hetcerina 

Dragon-flies and Damsel-flies 91 

clear-winged true damsel-flies. Most of them are small, and many keep 
so closely in low herbage or shrubby woodland that they attract little atten- 
tion. A few of the longer-bodied and longer-winged forms, however, fly 
in the open along the stream-banks or over the ponds. Some are strikingly 
varied with black and orange or yellow, and all, whether brightly colored 
09- dull, are graceful and charming. There are at least a dozen genera of 
Agrionids in this country, comprising about seventy-five species, but their 
classification is too difficult to be undertaken by general students. Damsel- 
flies deposit their eggs in the tissue of aquatic plants by cutting slits in the 
stems with their sharp ovipositor. The nymphs are slender and elongate, 
and can readily be known by the three caudal leaf-like tracheal gills. The 
nymph stage of these forms is much shorter than with the true dragon-flies, 
lasting usually probably but a few weeks, or at most two or three months. 
When ready to transform the nymphs crawl out of the water and into the 
low herbage on the stream or pond bank. I have seen scores of freshly 
emerged damsel-flies rising from a few square yards of tall grass near a pond, 
although it required close search to discover the nymphs, so well concealed 
were they in the dense tangle. 


Key to Families (Imagoes). 

Antecubitals of the first and second rows mostly meeting each other; triangle of 
fore wings with long axis at right angles to the length of the wings, triangle 
of hind wing with long axis in direction of the length of the wing. 


Antecubitals of the first and second rows not meeting (or running into each other) 
except the first and another thick one; triangles of fore and hind wings of 
similar shape (Fig. 121). 
Eyes meeting above on middle line of head; abdomen with lateral ridges. 


Eyes just touching at a single point or barely apart; abdomen without lateral 

ridges Cordulegasterid^. 

Eyes distinctly separated; abdomen without lateral ridges Gomphid^. 

Key to Families (Nymphs). 
Under-lip (labium) flat, not concealing most of the face, with jaw-like or oblong 
side pieces (lateral lobes). 

Antennae 7-segmented, tarsi 3-segmented, climbing nymphs. ..(Eschnid.e. 
AntenucB 4-segmented, the fourth segment rudimentary; fore tarsi 2-seg- 

mented; burrowing nymphs Gomphid^. 

Under-lip (labium) spoon-shaped, covering most of the face, when closed, with nearly 
triangular side pieces (lateral lobes). 

Two stout teeth with a notch between them on the middle lobe of the under- 
lip (labium) '. Cordulegasteridj!:. 

A single median tooth on the middle lobe of the under-lip LibelluliDjE. 

92 Dragon-flies and Damsel-flies 

The family Cordulegasteridae includes only seven species of dragon-flies 
found in the United States, all belonging to one genus, Cordulegaster. They 
are large, with eyes barely touching on top of the head, without metallic 
body-colors, and with clear wings. The nymphs burrow into the sand or 
vegetable silt on the bottom of shallow places. Thus buried, with only 
the top of the eyes and tip of the abdomen showing, they remain motionless 
for a long time, if prey does not come near. "In a dish of sand on my table," 
says Needham, "I have had a nymph remain without change of position 
for weeks, no food being offered it. Let any little insect walk or swim near 
the nymph's head, and a hidden labium springs from the sand with a mighty 
sweep and clutches it." The imagoes are strong flyers and have the habit 
of flying back and forth, as on a regular beat, over some small, clear stream. 

The family Gomphidae includes six genera, comprising about fifty species 
in our country. They are mostly large forms, clear-winged and with bodies 
striped with black and green or yellow. They are readily distinguished 
by the wide separation of the rather small eyes. The abdomen is stiff and 
spike-like. The eggs, held in a scanty envelope of gelatin, are deposited 
by the repeated descent of the flying female to the water of a clear pond 
or flowing stream, the tip of the abdomen first striking the surface. The 
gelatin dissolves and the eggs, scattering, sink to the bottom and become 
hidden in the silt. The nymphs are active burrowers, capturing their prey 
either on or beneath the surface of the bottom silt. The adults often alight 
on foliage, or on the surface of some log stretching across a stream, or on 
the bare soil of a path or roadway. They do not fly about in apparent 
sportiveness as the skimmers (Libellulidoe, p. 95) do, nor, like the skim- 
mers, perch atop a slender twig. June is the best month in the East for 
these dragon-flies. The principal genus of the family is Gomphus, which 
includes one-third of all our Gomphidae. Of these Gomphus exilis is 
probably the most common one in the Northeastern States. Its head is pale 
green, thorax brownish with two oblique green bands on each side, and 
abdomen blackish brown with a basal green spot or band on the back 
of each segment. The nymphs transform at the very edge of the water^ 
seldom crawling more than an inch or two above it. Hagenius brevistyliis 
is a large black-and-yellow species common in the East, South, and Middle 
West. The nymph has an unusually wide, flattened body. 

The /Eschnidse include our largest, swiftest, and most voracious dragon- 
flies. Various species are flying through the whole season from early spring 
to late summer. Some roam far from water, being found over dry fields 
and roadways, and even in houses. Some forms fly until late in the even- 
ing, making life a burden for the mosquitoes gathering for their night's 
singing and feasting. The eggs are thrust into the stems of aquatic plants, 
in floating timbers, in the wood of piers, etc., at or near the surface of the 

Dragon-flies and Damsel-flies 93 

water. The nymphs are slender, clean creatures, with smooth bodies pat- 
terned with green and brown, and very active, strong, and brave. They 
climb among green plants and roots or submerged driftwood along the border 
of open water or the edge of a current. The imagoes of this family can be 
recognized by the meeting of the eyes all along the top of the head. The 
wings are long, broad, and clear, and the body-colors are mostly bright blue 
and green. The family is represented in the United States by about twenty-five 
species, belonging to six genera. Anax Junius, one of the commonest dragon- 
flies all over the United States, and found also from Alaska to Costa Rica, 
in China, Siberia, and in various islands of the Pacific, notably the Hawaiian 
group, is the most inveterate enemy that the mosquito has. It is conspicu- 
ously on the wing from early spring to 
late fall, flying from daylight to dark, 
and doing untold good by its ceaseless 
warfare on the mosquito hosts. It 
can be recognized by its clear wings, 

large size (wings over two inches long), 

, , . , , , , , , Fig. 124a. Fig. 1240. 
and bright-green thorax and head, the 

, ^. , . ,1 r . Fig. 124a. — Top of head, showing charac- 

latter bearmg on the upper front a ^^^.^^ mark in front of eyes, of Anax 

round black spot surrounded by yellow, Junius. (Enlarged.) 

the yellow encircled by a dark-blue Fig. 1 24^. -Top of head, showing charac- 
•' •' teristic mark in front of eyes, of ALSckna 

ring (Fig. 124(7). A still larger member constricta. (Enlarged.) 

of this family is the great "hero" 

dragon-fly, Epiccschna heros, which is like Anax Junius in general appear- 
ance, but has wings two and one-half inches long, and abdomen nearly three 
inches long. It has a black T spot on the upper face, instead of a round 
one. Another similar, widely distributed and common form is .Eschna 
constricta, about the size of Afiax Junius, reddish brown marked with bright 
green, and with a black T spot on the upper front of face (Fig. 124^). The 
males have the abdomen marked with blue, with little or no green, while 
the females have but little blue or none at all. 

The members of the family Libellulidai are called "skimmers." They 
may be seen continually hovering over the surface of still water, or swiftly 
foraging over fields. Many of them have the wings strongly marked with 
large black or brown or milk-white blotches, and the abdomen is often 
covered with a whitish powder or "bloom." They outnumber all the other 
true dragon-flies in point of species, and except for Anax Junius, ^Eschna 
constricta, and perhaps the giant hero dragon-fly, include the most familiar 
and wide-spread members of the order. One of the best known and most 
beautiful of the skimmers is the pond-loving "ten-spot," Libellula pulchella 
(Fig. 125), found all over the country. Each of its wings has a longitudinal 
basal blotch, a median blotch (at the nodus), and an apical blotch of black- 


Dragon-flies and Damsel-flies 

ish brown. The males have the space between these blotches milky white. 
In old individuals the abdomen has a stron^ whitish bloom. Other familiar 

Fig. 125. — The ten-spot dragon-fly, Libellula pulchella. (After Needham; nat. size.) 

and well-marked species of Libellula are L. hasalis, with blackish-brown body 
and with the basal third to half of the wings dark brown or black and the 
rest of the wing clear, or in the old males chalky white out as far as the 

Fig. 126. — Libellula semi-Jasciala. (After Needham; natural size.) 

pterostigma, and in the females with brownish apices; L. qnadrimaculata, 
with olive or yellowish body marked with black, front wings with more 

Dragon-flies and Damsel-flies 


or less yellowish at base and along the front margin, and a small fuscous 
nodal spot, hind wings with a yellowish-black triangular basal spot and 
fuscous nodal spot; and L. semi-jasciata, whose complex wing-markings are 

Fig. 127. 

-The water-prince, Epicordnlia princeps, female. 
(After Needham; natural size.) 

shown in Fig. 126. Tramea is a genus of large swift dragon-flies whose 
hind wings have the base expanded and conspicuously colored. Tramea 
laceyata is a familiar species. The water-prince, Epicordulia princeps (Fig. 

5. — The amber wing, Perithemis domitia, male at left, female at right. 
(After Needham; natural size.) 

127), is a common large dragon-fly, but one hard to capture because of its 
fine flight. The wings show a basal patch, often nearly wanting on the 
front pair, a patch at the nodus, and a black apex. It likes "ponds or slug- 


Dragon-flies and Damsel-flies 

gish streams with muddy reed-grown banks, and seems absolutely tireless 
in flight; very rarely indeed is one seen resting." One of the smallest of 

Fig. 129. — The wind sprite. Celithemis eponina. (After Needham; natural size.) 

Fig. iT,o.—Tetragoneuria epinosa, female. (After Needham; natural size.) 

the true dragon-flies is the amber wing, Perithemis domitia (Fig. 128). The 
wings are clear amber, unmarked in the male, but richly spotted with dark 

Dragon-flies and Damsel-flies 


brown in the female. It has a slow hovering flight and often rests on the 
tips of erect reeds with wings held perfectly horizontal. It is only on wing 
in quiet, warm sunshine; clouds or cold breezes send them quickly into 
hiding. Among the familiar Libellulids with unblotched wings is Meso- 
themis simplicicollis, an abundant species east of the Rockies. The 
females and young males have head, thorax, and front half of abdomen 
green, the hinder half blackish brown. In old males the body becomes 
grayish blue with a whitish bloom. WiUiamson says that sometimes two 
males will flutter motionless, one a few inches in front of the other, when 
suddenly the rear one will rise and pass over the other, which at the same 
time moves in a curve downwards, backwards, and then upwards, so that 
the former position of the two is just reversed.*' These motions kept up 


Fig. 131. — The whitetail, Plathemis lydia. (After Needham; natural size.) 

with rapidity and regularity give the observer the impression of two inter- 
secting circles which roll along near the surface of the water. 

The whitetail, Plathemis lydia (Fig. 131), resembles the ten-spot, but 
is one-fourth smaller. In the males also the apex of the wings is usually 
clear, not brown. The whitetail rather likes slow-flowing brooks and 
open ditches. When alight it has the habit of setting its wings aslant down- 
ward and forward with a succession of jerks. Needham thinks that the 
powdery whiteness of the body of the old males (in females and young males 
the body is brown marked with yellow) must render it more easily seen by 
its enemies, the king-birds and others, and thus be a disadvantage in the 
struggle for existence. He says, indeed, that the whitest ones avoid rest- 


Dragon-flies and Damsel-flies 

ing-places over a dark background and settle oftenest on white sticks, on 
bleached stumps, or on light-colored earth. Very frequently one will alight 
on a white insect-net when it is laid down, or even when still held in the 



NCE when camping in the King's River 
Canon, one of the great vertical -walled, flat- 
floored canons of the Sierra Nevada, the 
boldest axeman of our party attacked the 
fallen trunk of a once towering yellow pine. 
The practical outcome of this attack was 
a sufficient supply of firewood for the 
cook's stone-built stove, but the great log 
yielded better things than chips and chunks. 
A few blows showed it to be the home of 
a thriving colony of the largest of the 
American termites {Termopsis angusticollis), and the thousands of indi- 
viduals in this insect household were objects of interested observation 
the summer through. We had heard of the rarity of white-ant queens in 
collections, and saw in this isolated and apparently easily "rounded-up" 
community an easy chance to discover the egg-laying queen of this species. 
But we had not reckoned with the Californ'a manner of tree-trunk: it 
outlasted the summer's chopping by two score feet of log four feet thick. 
Yellow pines grow 250 feet high in the Sierran forests. But although 
no queen was found, the make-up of the buried termite city was revealed. 
Galleries and chambers, secret ways and narrow tunnels were all ex- 
posed, and the interesting communal life of these soft, white-bodied little 
creatures was made partly known to us. 

We have in the United States but few kinds of termites, and these 
much less interesting in habit than those of tropic lands. The Amazons 
and Central Africa are the centers of termite life, and there, because of their 
great mounds, their serious ravages on all things wooden, and their enor- 
mous numbers, the white ants come to be nearly the most conspicuous of 
the insect class. Drummond's account, in his Tropical Africa, of the habits 
and life of the termites of the Central African region is simply a tale of 
marvels. And the scattered accounts of the Brazilian species are hardly 
less wonderful. In the South Sea, too, the termites play their part promi- 



The Termites, or White Ants 

nently. I have seen scores of cocoanut-palms in Samoa with their trunks 
traced over from ground to "feather-duster" top, a hundred feet above, by 
the laboriously builded wood-pulp tunnels of the termites. Each of these 
trees carried also on its trunk, about four feet from the ground, a termite 
"shed" or depot (Fig. 133), a foot thick, a foot wide, and two feet long, 
made, like the tunnels, of pellets of chewed wood, glued together with saliva, 
and filled with crowded galleries and chambers. 

Fig. 132.— Giant hillock-nests of termites in tropical Africa. 
(Adapted from Drummond.) 

But in the United States our few species make their communal nests 
in dead and dying wood, or underground, and not being given to building 
great dome-like mound-nests, or making covered ways up all the trees of 
a great forest or plantation, are not as conspicuous as their tropical cousins. 
Still, few observers of insects have failed to notice the Httle, white, wingless 
worker termites, scurrying about when some dead stump is overturned or 
split open, or to see the winged males and females swarming out of the 
ground some sunny day, and, after a brief period of flight, pursued by birds 
and predaceous insects, setthng to earth again and losing their wings. 

Before proceeding to take up the incompletely known life-history of our 
American termites, it will be advisable to describe their general structural 

The Termites, or White Ants 


characters and the composition of the termite communities. The body 
is always soft, and usually milky-whitish in color, though sometimes light 
or dark brown. It is plump, and slightly broader than thick. The abdo- 
men is joined broadly to the thorax, not by a little stem or peduncle as in 
the ants, with which insects the name "white 
ants" (unfortunately too long and widely 
used to be done away with) confuses the 
termites in the popular mind. The termites 
not only are not ants, but are neither nearly 
related to them nor of similar structure. 
The only resemblances between the two forms 
exist in the communal life and in the com- 
position of the community by different kinds 
ol' individuals. The termites are either blind 
or have only simple eyes, have slender an- 
tennae which look as if made up of tiny beads 
strung a-row, and have biting mouth-parts 
with strong jaws. They live in small or large 
communities, the individuals in any one of 
which, although belonging to the same species, 
being of from three to eight different kinds 
or castes. That is, each community is com- 
posed of winged and wingless individuals, 
the winged being males and females, while 
the wingless include immature individuals, 
sexually incomplete workers and soldiers, 
and also so-called complemental males and 
females which are individuals able to help 
in the increase of the community. In some 
species there are no workers, while in others 
the workers may be of two kinds. The 
soldiers differ from all the others in ihe 
extraordinary development of their jaws, 
which are long and scissor-like; their heads 
are also much enlarged and strongly chitin- 
ized. The food of all consists mainly of 
dead wood, and of curious pellets excreted 
from the intestine and called "proctodeal 

food." In addition some species attack live wood and even soft plants, 
and cloth, books, papers, etc., suffer from termite ravages. The serious 
nature of their attacks on wood will be referred to later. 

The development of the termites is apparently simple; the wingless 

Fig. 1.33. — Termite shed on 
cocoanut-palm in Samoa. From 
the shed note numerous tunnels 
leading down to the ground, in 
which is the main nest of the 
community; a few tunnels (only 
one visible in the picture) lead 
up the trunk of the tree. (Pho- 
tograph by the author.) 

I02 The Termites, or White Ants 

workers resemble closely, except in size, the just-hatched young; the soldiers 
have but to acquire their largeness of head and mandibles, and the perfect 
insects their wings. But there is a serious complexity in termite develop- 
ment in that at hatching all the young are alike, and the different castes 
or kinds of individuals become differentiated during the postembryonic 
development, i.e., after hatching. This matter is discussed later. 

In the United States but seven species of this order of insects are known. 
They represent three genera, which may be distinguished by the following 
table : 

Key to Genera. 

Simple eyes absent Termopsis. 

Simple eyes present. 

Tarsi with a pulvillus (little pad) between the claws; prothorax large and 

oblong; costal (anterior) area of the wings veined. .Calotermes. 

Tarsi without terminal pulvillus; prothorax cordate; costal area of wings 

without veins Termes. 

Termopsis and Calotermes each include two species, all four limited 
to the Pacific Coast; while Termes includes three species, of which but one, 
T. flavipes, is found in the northeastern states. This has been introduced from 
America into Europe, and is well known there. The other two species, and 
flavipes also, are found in the southwestern and Pacific coast states. Thus 
Termes flavipes (Figs. 134 and 135) is the only representative of the order Isop- 
tera which can be observed and studied in the East, 
but it is so commonly distributed that the student of 
insects in almost any locality can find its communities. 
Despite its abundance, however, the long time it has 
been known, and the very interesting nature of its 
habits, its life-history is not yet wholly worked out. 
Fig. 134. — T. flavi- Jt makes its nest in or under old logs and stumps. 

Marlatf natural Sometimes it mines a nest in the beams and rafters of 
size indicated by old houses. Howard records the serious injuries done 
^"^■■' to a handsome private residence in Baltimore through 

the mining of the first-floor timbers by the hidden termites. Comstock 
has found them in the southern states infesting Uving plants, particularly 
orange-trees, guava-bushes, and sugar-cane. According to Comstock, they 
attack that part of the living plant which is at or just below the surface 
of the ground. In the case of pampas-grass the base of the stalk is 
hollowed; with woody plants, as orange-trees and guava-bushes, the bark 
of the base of the trunk is eaten, and frequently the tree is completely 
girdled; with sugar-cane the most serious injury is the destruction of the 

The Termites, or White Ants 


The workers of T. flavipes (Fig. 134) are, when full grown, about ^ in, 
long, while the soldiers are a little larger. Both of these castes are whitish. 
But the winged males (Fig. 135c) and females which come from the nest 
and swarm in the air in late spring or early summer are chestnut-brown 
to blackish and measure about I in. in length. The four wings are of about 
equal size, and when the insect is in flight expand about f in. When at 
rest they lie lengthwise on the back, projecting beyond the tip of the abdo- 
men. They have many veins and are pale brown in color. After fl3'ing 
some time and to some distance, the insects alight on the ground and shed 
their wings (Fig. 1356). This they are enabled to do because of a curious 
suture or line of weakness running across each wing near its base. All the 
wing beyond this suture falls oS, leaving each now wingless male or female 
with four short wing-stumps. These swarming flights 
attract the birds. Hagen noted fifteen different species 
of birds following such a termite flight one May-day in 
Cambridge, Mass. "Besides the common robins, blue- 
birds, and sparrows," he says, 
"were others not seen before 
near the house. The birds 
caught the Termes partly in 
flight, partly on the ground, 
and the robins were finally 
so gorged in appearance that 
their bills stood open! " 

After the swarming flight 
the few uneaten males and 
females pair, and each pair 
probably founds a new colony. 
Perhaps some of the pairs 
are found by workers, and 

taken possession of as the royal couple for a new community. Exactly 
how the new communities of flavipes begin is not known; and this is 
an excellent opportunity for some amateur observer to distinguish himself! 
The egg-laying queen mother of a flavipes colony also has yet to be 
discovered. There exist in many species of termites individuals caUed com- 
plemental males and females. These are forms which, in case of the loss 
of the real king or queea, can develop into substitute royalties. Whether such 
forms exist in all flavipes colonies does not seem to be certainly known. 
It is obvious that there is still much to learn about the interesting life of 
our commonest and most wide-spread termite species. 

Of the other six species of our country, afl of which are limited to the 
southern, southwestern, and Pacific states, three, representing all of the 

Fig. 135a. Fig. 135&. 

Fig. 135a. — T. flavipes, winged male. (After Mar- 

latt; natural size indicated by line.) 
Fig. 135&. — T. flavipes, complementary queen. 

(After Marlatt; natural size indicated by line.) 


The Termites, or White Ants 

three genera, and found about Stanford University, have been recently 
studied by Professor Harold Heath. These are Termopsis angusticollis, 
the largest of the American termites, Calotermes castaneiis, a small species with 
brown-bodied winged forms, and Termes lucijugus, a small white species 
common in Europe, and probably brought to this country from there. The fol- 
lowing account of Termopsis angusticollis is based chiefly on Heath's * studies. 
Termopsis angusticollis (Fig. 136) is the largest of the three species and 
the most abundant. In favorable localities colonies may be found in almost 
every stump and decaying log, and even in dead branches on otherwise healthy 
trees. The galleries are made in the deeper portions of the wood, and 
usually follow the grain. The colonies with the primary royal pair number 
usually from 50 to 1000 individuals, and include workers, soldiers, and im- 
mature forms. The full-grown workers (Fig. 136) are f in. long, the soldiers 
(Fig. 136) § in., and the kings and queen (Fig. 137) a little less, while the 
wings expand i\ in. After the death of the primary royalties and the 
development of several substitute royal ^ 

forms the egg-laying and consequent 
increase of the colony are much more 
rapid. Heath counted 3221 individuals 

Fig. 136. Fig. 157. 

Fig. 136. — The large termite of California, Termopsis angusticollis; workers, young, 

and a soldier. (From life; natural size.) 
Fig. 137. — A, Dealated primary queen of Termopsis angusticollis, at least four years 

old; B, complemental queen. (After Heath; three times natural size.) 

in one colony, in which were also thousands of eggs. The colony which we 
found in the yellow-pine log in the King's River Cafion certainly num- 
bered many thousands. In the late summer or early autumn the nymphs 
(young stage, with visible wing-pads of perfect insects) that have developed 
during the year moult, the operation taking from ten to twenty minutes, 
after which they rest for two hours, while the wings expand, and the 
body-wall hardens and darkens; they take flight usually about dusk. Some 

* Heath, H., The Habits of California Termites, Biological Bulletin, vol. 4, 1902, 
PP- 47-63. 

The Termites, or White Ants 105 

soon fall to the ground, but others may fly a mile. The swarm is pursued 
by birds until dark, and then bats take a turn at the chase. The few ter- 
mites that escape fly from tree to tree, seeking a spot of decaying wood. 
Heath has noted them dashing against door-knobs and nail-holes and against 
discolored spots on trees and logs, in their search for a place where decay 
has begun. After finding a suitable spot they usually shed their wings, 
not by biting them off, as said of some species, but by curving the abdomen 
until it rests across the wings of one side and then moving backwards 
and sidewise until the wing tips are brought against some obstruction, 
thus causing the wings to buckle and break along the transverse suture or 
line of weakness at the base. Sometimes the wings are not shed until after 
the nest is begun. The spot is usually selected by the female, and she begins 
the mining and does most of it. She is accompanied by one or more males, 
who may occasionally help in excavating. When the burrow is large enough 
for two, one male usually crowds in beside the queen and fights off the others. 
Sometimes two males may remain with the queen; Heath thinks that such 
a condition may last for a year or more. He has found a few cases where 
two, three, and even six pairs live in company. The actual mating does 
not take place, probably, until some time after the nest is begun. Heath 
has notecT pairing from a week to a fortnight after swarming. 

The egg-laying may be long postponed. Usually, however, about two 
weeks after pairing the first egg is laid, and from one to six are deposited 
daily until the total number amounts to from fifteen to thirty. When the 
habitat is unusually moist the royal pair may remain together for a year 
without producing young. Heath has found the Termopsis royalties to 
mate readily in captivity, and has had more than 500 pairs of primary kings 
and queens in excellent condition after a year of captivity. Royal pairs 
with small colonies are readily found by stripping oft' the bark of trees from 
three to nine months after the swarming period. Heath has been the first 
to find actual egg-laying queen termites in this country. 

After from fifteen to thirty eggs are laid the laying ceases, and the 
parents give their time to enlarging the nest and to caring for the eggs, 
which are kept scrupulously clean, and frequently shifted from place to 
place in the nest. The young are all alike when first hatched. After three 
moults, one of them appears as a large-headed individual, and after three 
more moults develops into a perfectly formed soldier, although little more 
than one-half the size of the soldiers in old communities. Three months 
later another soldier appears, larger than the first, and later others still 
larger, until after a year the full-sized form appears. The first workers, 
too, are smaller than the later ones. Nymphs, i.e., young of the winged 
individuals, do not appear until after the first year, so that the swarm of 
winged individuals cannot leave a nest until the end of the second year of 


The Termites, or White Ants 

its existence. The life of these early, undersized individuals is short. 
They disappear, perhaps are killed, when the full-sized individuals appear. 
These latter, both workers and soldiers, live at least two years and perhaps 

The primary king and queen hve for at least two years, and almost cer- 
tainly longer. Heath believes he has evidence of five years of life. After 
the death of the royal pair from natural or other causes, the members of 
the orphaned colony develop from the young nymphs from ten to forty sub- 
stitute royal forms. By some unknown process, perhaps peculiar feeding, 
these selected nymphs are quickly brought to sexual maturity, and the queens 
begin egg-laying. As they are fed and cleaned by the workers, their only 
business is to lay eggs. Heath observed some of the larger queens to lay 
from seven to twelve eggs a day continuously. In exceptional cases a 
worker, or even a soldier, may be developed into an egg-laying queen. 
One may also occasionally find a few winged soldiers. 

In Africa forty-nine species of Termites are known * (Sjostedt), and it is 
on this continent that "the results of Termitid economy have reached their 
climax." More than a century ago an exploring Englishman, Smeathman, 
startled zoologists with his account of the marvelous termite communities 
of West Africa. He told of the great mound- 
nests of Termes heUicosus, twenty feet high, and 
so numerous that they had the appearance of 
native villages (Fig. 132). The soldiers are fifteen 
times as large as the workers, and the fertile 
queen has her abdomen so enlarged and stretched 
by the thousands of eggs forming inside that it 
comes to be "fifteen hundred or two thousand 
times the bulk of the rest of her body and 
twenty or thirty thousand times the bulk of a la- 
borer." He describes the egg-laying as proceed- 
ing at the rate "of sixty a minute, or eighty thou- 
sand and upward in one day of twenty-four 
hours." In the South Kensington Museum at 
London there is such a prodigious queen resem- 
bling simply a cylindrical whitish sausage four 
inches long. A similar specimen is to be found 
in the natural-history museum of the University 
of Kansas. 
The enormous number of individuals in a great village of nests cannot 

Fig. 138. — Worker and 
queen of Termes red- 
mani. (After Nassonow; 
natural size.) 

* Sjostedt, Y., Monographie der Termiten Afrikas, Kongl. Svenska, Vetensk. Ak. 
Handl., v. 34, 1900, pp. 1-236, Stockholm. 

The Termites, or White Ants 107 

even be imagined. But according to African travelers the direct results 
of the presence of such a population are very apparent. Drummond 
(Tropical Africa, 1891) writes: "You build your house, perhaps, and for 
a few months fancy you have pitched upon the one solitary site in the coun- 
try where there are no white ants. But one day suddenly the door-post 
totters, and lintel and rafters come down together with a crash. You look 
at a section of the wrecked timbers, and discover that the whole inside is 
eaten clean away. The apparently solid logs of which the rest of the house 
is built are now mere cylinders of bark, and through the thickest of them 
you could push your little finger. Furniture, tables, chairs, chests of drawers, 
everything made of wood, is inevitably attacked, and in a single night a 
strong trunk is often riddled through and through, and turned into match- 
wood. There is no limit, in fact, to the depredation of these insects, and 
they will eat books, or leather, or cloth, or anything; and in many parts of 
Africa I believe if a man lay down to sleep with a wooden leg it would be 
a heap of sawdust in the morning! So much feared is this insect now that 
no one in certain parts of India and Africa ever attempts to travel with such 
a thing as a wooden trunk. On the Tanganyika plateau I have camped on 
ground which was as hard as adamant, and as innocent of white ants appar- 
ently as the pavement of St. Paul's, and awakened next morning to find a 
stout wooden box almost gnawed to pieces. Leather portmanteaus share 
the same fate, and the only substances which seem to defy the marauders 
are iron and tin." 

But more impressive than this devastation of houses, tables, and boxes is the 
sight of millions of trees in some districts plastered over with tubes, galleries, 
and chambers of earth due to the amazing toil of the termites in their search 
for dead or dying wood for food. According to Drummond, these tunnels 
are made of pellets of soil brought from underground, and stuck together 
with sahva. The quantity of soil thus brought above ground is enormous 
and Drummond sees in this phenomenon a result very similar to that accom- 
plished by earthworms in other parts of the world, and made familiar to 
us by Darwin, namely, a natural tillage of the soil. As Drummond says: 
"Instead of an upper crust, moistened to a paste by the autumn rains and 
then baked hard as adamant in the sun, and an under soil hermetically sealed 
from the air and light, and inaccessible to all the natural manures derived 
from the decomposition of organic matters — these two layers being eter- 
nally fixed in their relations to one another — we have a slow and continued 
transference of the layers always taking place. Not only to cover their 
depredations, but to dispose of the earth excavated from the under- 
ground galleries, the termites are constantly transporting the deeper 
and exhausted soils to the surface. Thus there is, so to speak, a con- 
stant circulation of earth in the tropics, a ploughing and harrowing, not 

io8 The Termites, or White Ants 

furrow by furrow and clod by clod, but pellet by pellet and grain by 

With a few references to certain special conditions and problems in the 
termite economy, we must finish our consideration of these highly inter- 
esting insects. Do the termite individuals of a community communicate 
with each other, or is the whole life of the colony so inexorably ruled by 
instinct that each individual works out its part without personal reference 
to any other individual, although with actual reference to all the others, 
that is, to the community as a whole? It is pretty certain that termites have 
a means of communication by sounds. The existence of a tympanal audi- 
tory organ in the tibiie of the front leg, like that of the crickets and katy- 
dids, has been shown by Fritz Miiller, and the individuals have a peculiar 
jerking motion which seems likely to be connected with the making of 
sounds by stridulation, sounds, however, that are not audible to us. 

The spread of termites from one continent to another, as in the case of 
Termes flavipes from America to Europe, and Termes lucijugus from 
Europe to America, can be easily explained by involuntary migration in 
ships. In unpacking several cases of chemicals received from Ger- 
many at Stanford University, scores of termites were exposed when the 
wooden boxes were broken up. The insects, mining in the wood of the 
boxes, had protection, food, and free transportation on their long ocean 
journey from Hamburg around Cape Horn to California! 

In termite nests are often found individuals of other insect orders. So 
often are such cases noted, and so many are the kinds of strangers likely 
to be present, that entomologists recognize a special sort of insect economy 
which they term termitophily, or love of termites! The strangers seem to 
be tolerated by the termites, and apparently live as guests or conmensals. 
More than loo species of insects have been recorded as termitophiles. This 
curious guest-life exists on even a much larger scale in the nests of true 
ants, in which connection it is called myrmecophily (see p. 552), 

The most important problem, and one whose solution will require much 
exact observation (and, if possible, experimentation), is that of the origin, 
or causes of production, of the different castes or kinds of individuals in 
the termite community. It has been determined by various observers that 
all the termites of a community are apparently alike at birth. That is, 
there is no apparent distinction of caste, no separation into workers, soldiers, 
and perfect insects. The soldiers and workers are not, as was formerly 
thought, the result of the arrested development of the reproductive organs. 
They are not restricted to one of the sexes. If then it is not arrested develop- 
ment, or sex, or embryonic (hereditary) differentiation, what is the causal 
factor? Grassi, an Italian student of the termites, thinks that it is food; 
that the feeding of the young with food variable in character brings about 

The Termites, or White Ants 109 

the differentiation of individuals. To understand this claim it is necessary 
to attend more closely to the feeding habits. The food of termites con- 
sists almost exclusively, as has already been said, of wood. But this wood 
may be. taken directly from the walls of the burrow or secured indirectly 
from another individual. In this latter case it consists of disjecta of undi- 
gested material, which, while mostly wood, must be mixed with other or- 
ganic material: because the termites keep their nests clean by eating their 
cast skins and the dead bodies of other individuals. This undigested mate- 
rial is called proctodeal food. In addition, a certain amount of evidently 
very different matter is regurgitated through the mouth from the anterior 
part of the alimentary canal. This is called stomoda^al food. As the young 
receive all their food from the workers, it is apparent that there is oppor- 
tunity for a choice, on the part of the nurses, in the kind of food given the 
young. And it is presumed by Grassi that such a choice is made, and that 
it results in the differentiation of the castes. As a matter of fact, such a 
differentiation of individuals is accomplished in the honey-bee community 
by feeding those larva? which the workers wish to make fertile queens "royal 
jelly" — a rich food regurgitated through the mouth from the anterior part 
of the alimentary canal. This is done for the queens during the whole 
larval life, while larvae which are fed royal jelly for only one or two days, 
and then mixed pollen and honey for the rest of larval life, develop into 
workers. With the honey-bee, however, the workers are to be looked on 
as probably only arrested females. But in the case of Termopsis angusti- 
coUis Heath has experimented by feeding members of 
various colonies, both with and without primary royal 
pairs, "on various kinds and amounts of food — procto- 
deal food dissected from workers, or in other cases from 
royal forms, stomodeal food from the same sources, 
sawdust to which different nutritious ingredients had 
been added — but in spite of all I cannot," he says, 
"feel perfectly sure that I have influenced in any un- 
usual way the growth of a single individual." 

This is by all odds the most important and interesting 
problem in termite economy, and the solver of it will do 
much for zoological science. 

A singular and primitive family of small insects, the p ^ , . 

Embiida?, of doubtful affinities, is represented by not iexana. (After 
more than twenty hving species, of which but four Melander; en- 
occur in this country. The individuals do not live in 
communities as the termites do, but in structural characters they probably 
more nearly resemble these insects than any others. Fig. 139 illus- 
trates a typical Embiid. This species, Embia iexana, is about one-quarter 

1 1 o The Termites, or White Ants 

of an inch long, and of rufous color. It was described from a few specimens 
found at Austin, Texas. This insect seems to be very susceptible to differ- 
ing degrees of humidity, and specimens were visible only after the ground 
had been moistened by rains. As the sun dries the ground, the insects 
burrow into the soil. They spin small silken webs in which they live singly. 
These webs are tunnels made in some crevice of the rock which shelters 
them, or are spun between grains of soil. They are an inch or more in 
length and closed at one end, and probably serve simply for protection. The 
spinning-organs of the insect are located in its fore feet, a condition unique 
among animals. The food-habits of the Embiids are not yet known. 


THE BOOK-LICE AND BARK-LICE (Order Corrodentia) 
AND THE BITING BIRD-LICE (Order Mallophaga) 

OMETIMES in taking from the shelves an old 
book, long untouched, there may be seen, on 
turning its leaves, numerous extremely minute, 
pale-colored, wingless insects, the book-hce, or 
dust-hce. So small are they, indeed, that a 
reading-glass or hand-lens will be needed to make 
out anything of their real appearance. They 
run about rather swiftly and seek to conceal their soft, defenceless little 
bodies somewhere in the binding. Under the lens they are seen to have 
a rather broad, flattened body (Fig. 140), six short legs, no wings (although 
sometimes tiny wing-pads are present), long, slender antennae, and a pair of 
small black spots on the head, the simple eyes. There is a distinct neck, 
the head being free, and plainly wider than the prothorax. The abdomen 
is nearly oval in outline. There are no distinctive markings or pronounced 
chitinization of the soft body-wall. These book-lice can be found else- 
where than in old books; they feed on dry, dead organic matter, the 
paste of the book-bindings and the paper, and are common in birds' nests, 
where they feed on the cast-off feathers, in the crevices of bark, and on 
old splintered fences, where they feed on moulds and dead lichens. 

Certain other insects closely related to the book-lice are not so small and 
simple, however, some having two pairs of wings and a plump, rounded 
body (Fig. 141); these look much like plant-lice (Aphids). These winged 
kinds do not live in libraries, moreover, and the name "book-lice" is a 
misnomer for them. They are rarely seen by persons not trained entomolo- 
gists, and indeed are not at all familiar to professed students of insects. 

The life-history of these obscure insects has been but little studied, but it 
is of a simple kind, the metamorphosis being incomplete, and in the case 
of the wingless forms certainly very slight. The young of the wingless forms 
" greatly resemble the old, but have no ocelli or wings, and sometimes the 
tarsi are of two joints, while in the adult they have three." The structure 
of the adults presents no points of particular interest except in the case of 
the mouth. The book-lice have biting mouth-parts, the jaws being strong 
and heavy for the successful mastication of the hard dry food. In the throat 

1 1 2 Book-lice and Bark-lice ; Biting Bird-lice 

there is a peculiar little chitinized structure, which may be called the 
oesophageal sclerite (Fig. 145). This structure is situated in the floor of 
the pharynx (forward end of the oesophagus), and has some special function 
in connection wfth the peculiar food-habits. It was first described by Bur- 
gess, and was for a long time supposed to be peculiar 
''J^T } to the book-lice alone. But, in a study of the mouth 
structure of the biting bird-lice (Mallophaga) , I found 
an almost identical oesophageal sclerite in thirteen out 
of the twenty-two genera of the Mallophaga. On 
. ,, the basis of this common possession of a curious 
and undoubtedly important mouth structure by the 
book-lice and the bird-lice (and on the basis of other 
strong similarities) it seems certain that these two 
groups of insects have a common ancestry not very 
remote, and probably should be included in a single 
Fig. 140. — A wingless order, 
book -louse, Atropos sp. -pj^g Q^der Corrodentia as at present known con- 
tains about two hundred described species, scattered 
over the world. The largest species occur in Brazil, and have an ex- 
panse of wing of nearly an inch. Ceylon and the Hawaiian Islands are 
said by Sharp to be. specially rich in species. 

The members of the order can be divided into two families as follows: 

Wings well developed; ocelli present (in addition to compound eyes). . .PSOCID.E. 
Wings wanting or present as small scales or pads; ocelli absent . — Atropid.^. 

The winged Corrodentia or Psocidae (which may be called bark-lice to 
distinguish them from the wingless book-lice) are ^ 

too rarely seen to be at all familiar. They may /^-^^^ 

most commonly be found in small clusters on bark, ^-' 

each cluster or colony being covered over by fine 
silken threads spun from the mouth. The wings 
are held roof-shape over the back (Fig. 141), and 

the body and wings are usually pale smoky in ^^^ i4i._a winged 
general color. The small white eggs are laid on the bark-louse. (Thirteen 
surface of the bark in small patches, and in a cluster ^^™^^ '^^^"'"^^ ^'^^-^ 
of bark-lice, individual in all stages, from very young to adult, may be seen. 

Banks gives the following key to the North American genera: 

The techinal terms discoidal cell and posterior cell may be understood by reference to 
Fig. 142. 

1. Wings with scales and long hairs Amphientomum. 

Wings without hairs and scales, hyaline 2. 

2. Tarsi 3-jointed 3. 

Tarsi 2-jointed 4- 

Book-lice and Bark-lice; Biting Bird-lice 113 

3. Discoidal cell closed Myopsocus. 

Discoidal cell open , .' Elipsocus. 

4. Discoidal cell closed 5. 

Discoidal cell open 6. 2 

5. Discoidal cell four-sided. Psocus. t ^.<;)>i'''ZI^^^^^--'^ " ^^ >^\ «?" 

Discoidal cell five-sided. 


6. Third posterior cell elliptical. 


Third posterior cell elongated. 

PoLYPSOCUS ^'*^' ^4^' — Diagram of venation of a Psocid. 
„, . , , . 11 . , ' d, discoidal cell; \a, 2a, -la, posterior cells. 

Third posterior cell absent. ^^^^^^ g^^^^ -, 

The few North American species of the true book-lice or Atropidae are 
included in five genera, which may be distinguished as follows: 

The technical terms, hitherto undefined, used in the following table are the following: 
squaincp, wings in the condition of small scales or pads; hyaline, clear, not colored. 

1. Meso- and metathorax united, no wings Atropos. 

Meso- and metathorax separate, rudimentary wings 2. 

2. Wings with veins Dorypteryx. 

Wings veinless, in form of squamte or tubercles 3. 

3. Squamas small, hyaline Clothilla. 

Squamae in the form of scars Lepinotus. 

Small tubercles in the place of squamas Hyperetes. 

The genera Atropos and Clothilla were named for two of the three Fates 
of mythology, and a third genus was named Lachesis for the third Fate, but 
unfortunately the last genus was not a valid one, so the book-lice have lost 
their third Fate, and by the rigid laws of zoological nomenclature can never 
regain her! The few species of these two Fate-named genera are the com- 
monest of the book-lice. Atropos divinatoria is the species usually 
found in books. It is about i mm. long, is grayish-white, and the small 
eyes show as distinct black specks on the head. It does not limit its feeding 
to the paste of book-bindings, but does much damage to dried insects in 
collections. To this insect has long been attributed the power of producing 
a ticking noise known as the "death-watch," but McLachlan, an authority 
on the Corrodentia, does not believe that this minute insect "with a body 
so soft that the least touch annihilates it can in any way produce a noise 
sensible to human ears." A small beetle, called Anobium, is well known 
to make such a ticking (by knocking its head against the wood of door-casings, 
floors, etc., in which it lives) and probably the "death-watch" is always 
made by this beetle. 

Bird-collectors are often annoyed by small, wingless, flat-bodied, swift- 
running insects which sometimes escape from the feathers of bird specimens 
to the hands and arms of the collector. Poultry-raisers are sometimes more 
seriously troubled by finding them so abundant on their fowls as to do con- 

114 Book-lice and Bark-lice; Biting Bird-lice 

siderable injury. They are called bird-lice, but they should not be confused, 
because of this name, with the true blood-sucking lice that infest many kinds 
of animals, particularly domestic mammals and uncleanly persons. The 
biting bird-lice (Fig. 143), constituting the order Mallophaga, never suck 
blood, but feed exclusively on bits of the dry feathers, which they bite off 
with small but strong and sharp-edged mandibles. The true lice have 
mouth-parts fitted for piercing and sucking, and 
constitute one of the numerous families of the 
order of sucking bugs, Hemiptera (see p. 217). 

More than a thousand species of biting bird- 
lice, or Mallophaga, are known, of which about 
two hundred and fifty have been found on North 
American birds. Although by far the larger num- 
ber of Mallophaga infest birds, numerous species 
are found on mammals. On these hosts the insect 
feeds on the hair or on epidermal scales. On 
both birds and mammals, therefore, the food con- 
sists of dry and nearly or quite dead cuticular sub- 
stances, and never of blood or live flesh. Those 
species of Mallophaga which infest birds are never 
found on mammals, and vice versa. 

The injury done to the hosts by these parasites 
consists not in the character of the food-habits, but 
chiefly in the irritation of the skin caused by the 
scratching of the sharp-clawed little feet of the insects 
in their migrations over the body. When, as hap- 
pens sometimes in poultry-yards and dovecotes, 
a fowl or pigeon is infested by hundreds of these active little pests, the 
afflicted bird becomes so restless and excited that it takes too little food 
and gets too little rest and thus grows thin and weak. The dust-baths 
taken by fowls and other birds are chiefly to get rid of the bird-lice. The 
fine dust, getting into the breathing-pores (spiracles) of the insects, suffocates 
them. So that the best remedies for these pests of the barn-yard are to 
see that the fowls have plenty of dust to bathe in, and also to keep 
thoroughly clean their roosting- and breeding-places. By tightly closing 
up the hen-house and burning sulphur inside (the fowls, it is hardly necessary 
to say, first being excluded) most of the infesting parasites can be killed. 

The life-history of the Mallophaga is very simple. The small elongate 
eggs are glued separately to the hair or feathers of the host, and from them 
young soon hatch (Fig. 144,3), which, except in size and, to some degree, in 
marking, closely resemble the parents. These young begin immediately their 
hair or feather diet, grow larger, moult a few times, and in a few weeks reach 

Fig. 143. — A biting bird- 
louse, Nirmiis prce- 
stans, from a tern, 
Sterna maxima. (Pho- 
tomicrograph by 
George E. Mitchell; 
natural size, one-twelfth 

Book-lice and Bark-lice; Biting Bird-lice 115 

maturity. There is never, in young or old, any sign of wings or wing-pads. 
The body is flattened, so much so indeed that it is difficult to hold a live 
specimen securely between thumb and finger-tip. The body-wall is strongly 
chitinized, and is firm and smooth. The markings are often very distinct, 
and sometimes bizarre, but the coloration varies only from white to black 
through various shades of pale yellowish brown, tawny, reddish brown, and 
blackish brown. The antennae are short and in one suborder (see classifica- 
tion key) are wholly concealed in pits or grooves on the under side of the 

Fig. 144. — Immature and adult stages of the biting bird-louse, Lipeuriis forficidatiis, 
taken from a pelican, i, adult female; 2, adult male; 3, very young stage; 4, 
older immature stage. (Natural size of adult specimens ^s in.) 

flattened head. The legs are strong, and each foot bears two claws. These 
small creatures run very swiftly. 

Perhaps the oddest thing about the structure of the Mallophaga is the 
presence in the throat of the curious oesophageal or pharyngeal sclerite 
already referred to in the account of the Corrodentia. This sclerite is a 
sort of bonnet-shaped piece (Fig. 145) lying in the lower wall of the throat 
and seems to be an arrangement for starting the little bitten-off pieces of 
feather barbs straight, that is, lengthwise down the oesophagus! The bark- 
lice and book-lice, which have a similar oesophageal sclerite, also bite off 
and swallow small bits of hard, dry organic substance. 

I 1 6 Book-lice and Bark-lice ; Biting Bird-lice 

The most interesting thing in connection with the Mallophaga, excepting 
their parasitic life and strange food-habits, is the puzzHng problem of their 
distribution. The problem in its largest phase is this: the species of Mal- 
lophaga are, in a majority of cases, peculiar (so far as recorded) each to some 
one host species. But the instances are many where a single parasite species 
is common to a few or even to many host species. How does this condition 
of commonness to several hosts come to exist? 

As the Mallophaga are wingless, their power of migration from bird to 
bird is limited. Moreover, they can live for but a short time oiT the body 

of a warm-blooded host. After a bird is 
shot, the Mallophaga on it die in from 
two hours to three or four days: in rare 
cases living individuals are found on the 
drying bird -skin after a week. Although 
the parasites in a badly infested hen-house 
will be seen on the roosts and in the nests, 
in Nature the insects are rarely found off 
the host's body. On such a likely place 
as an ocean rock from which I had just 
frightened hundreds of perching pelicans, 
cormorants, and gulls no parasites could 
be found. Practically migration must be 
accomplished while the bodies of the hosts 

sclente, lateral aspect, from a biting _ . 

bird-louse, Euryjnopetiis taurus, from mating and nesting, and when gregarious 
an albatross. (Greatly magnified.) |^jj.^g ^^^^^ ^^ p^j-^j^ closely together. 

Occasional migration might occur from a bird of prey to its captured 
victim, or from victim to hawk. 

The general character of the cases in which a single Mallophagan species 
is common to several host-species may now be considered. Docophoriis 
lari has been found on thirteen species of sea-gulls, and Nirmiis lineolatiis 
on nine. Gulls are gregarious, perching " together on large food-masses 
and on ocean rocks. But on these rocks gulls are closely associated with 
other coast birds, as cormorants, pelicans, murres, etc. And the gull-para- 
sites might have opportunities to migrate to these other bird-species. 
Docophorus iderodes and Trinoton luridum are common to many duck species 
(each has been collected from nine), but ducks also are gregarious, and in 
addition are much given to hybridizing. But a parasite may be common to 
several host-species of non-gregarious habits. Docophorus platystomiis is 
common to several hawk-species, D. cursor to several owl-species, D. excisus 
to several swallows, D. californiensis to several woodpeckers, and Z). com- 
munis to several passerine birds. In the other genera of Mallophaga are 


-Bonnet-shaped pharyngeal 

Book-lice and Bark-lice; Biting Bird-lice 117 

similar cases, and in all these cases it is hard to see how actual migration 
of the parasite from host to host of different species could take place. Indeed 
there are cases in which such migration is absolutely impossible. Of the 
262 species of Mallophaga taken from North American birds, 157 have 
been described as new species, while 105 are specifically identical with Mal- 
lophaga originally described from European and Asiatic birds; hosts, that 
is, not only of different species, but geographically widely separated from 
the North American hosts! Eliminating the few cases of importations of 
Hving European birds to this country, and the few species of cicumpolar 
range, there remain to be accounted for about 100 cases in which a single 
species of Mallophaga is common to both Old World and New World hosts. 

It will have been noted that in all the cases above mentioned of parasite 
species common to several North American host-species, the host-birds are 
closely allied forms, that is, species of the same genus or allied genera. 
This condition holds good also for practically all of the cases in which both 
European and American hosts have a common parasite. For example, 
Docophorns pertusus is common to the European coot {Fulica atra) and 
the American coot {Fulica americana); Nirmus pileus is common to the 
European avocet {Reciirvirostra avocetta), and to the American avocet 
{Recurvirostra amcricana) ; Lipeurus jorficulatus is common to the European 
pelican {Pelecanus onocrotalus) and to the American pelicans (Pelecanus 
erythrorhynchus and P. calif ornicus), and so on through the list. From 
this fact of near relationship of hosts in all the cases of parasite species com- 
mon to several host-species it seems almost certain that this common occur- 
rence, under circumstances not admitting of migration of the parasites from 
host to host, is due to the persistence of the parasite species unchanged from 
the time of the common ancestor of the two or more now distinct but closely 
allied bird-species. In ancient times geographical races arose within the limits 
of the ancestral host-species; these races or varieties have now come to be dis- 
tinct species, distinguished by superficial differences in color and mark- 
ings of plumage, etc. But the parasites of the ancient hosts have remained 
unchanged; the plumage as food, the temperature of the body, practically 
the whole environment of the insect, have remained the same; there has 
been no external factor at work tending to modify the parasite species, and 
it exists to-day in its ancient form, common to the newly arisen descendants 
of the ancient host. 

To classify Mallophaga the following keys to suborders, families, and 
genera may be used. In these keys are included only genera which have 
been found in the United States. Seven other genera of Mallophaga are 

In the following tables the following technical terms are used which have not been 
previously defined: clavate, club-shaped; capitate, with the tip swollen like a ball;» tra- 

1 1 8 Book-lice and Bark-lice ; Biting Bird-lice 

hecula, triangular membranous processes projecting laterally from the head and situated 
one in front of each antenna; temples, the hinder lateral parts of the head; ocular emar- 
gination, a bending in of the lateral margins of the head just in front of the eyes; lahral 
lobes, short blunt membranous processes projecting laterally from near the front angles 
of the head; sternal markings, blackish markings, bars or spots, on the ventral aspect of 
the thorax. 

Key to Suborders of Mallophaga. 

With short slender 3- or 5 -segmented, exposed antennae; no palpi; mandibles 

vertical Ischnocera. 

With short clavate or capitate, 4-segmented antennae concealed in shallow cavities 
on under side of head; 4-segmented palpi; mandibles horizontal. 

Key to Genera of the Suborder Ischnocera. 
A. With 3-segmented antennae; tarsi with one claw; infesting mammals only (family 

TrichodectidcE) Trichodectes. 

AA. With 5-segmented antennae; tarsi with two claws; infesting birds only (family 
B. Antennas alike in both sexes. 

C. Front deeply angularly notched Akidoproctus. 

CC. Front convex, truncate, and rarely with a curving emargination, but never 
angularly notched. 
D. Body broad and short; head with large movable trabeculae. 

E. Forehead with a broad, transverse membranous flap, project- 
ing beyond lateral margins of the head in the male, barely 

projecting in the female Giebelia. 

EE. Without such membranous flap Docophorus. 

DD. Body elongate, narrow; head with vers'- small or no trabeculae. 


BB. Antennae differing in the two sexes. 

C. Body wide, elongate oval to suborbicular. 

D. Temples rounded; tip of abdomen with shallow, curving emargina- 
tion; antennae with no appendage; third segment unusually long. 


DD. Temples usually angulated; tip of abdomen convex, rarely angularly 
emarginated with two points. 

E. First antennal segment of male large, and sometimes with 
an appendage; third segment always with appendage. 

EE. First antennal segment of male large, but always without 
appendage; third segment without appendage. .Goniocotes. 
CC. Body elongate, narrow, sides subparallel. 

D. Antennae and legs long; a semicircular depression in front of 

moutli LiPEURUS. 

DD. Antennae and legs short; depression in front of mouth narrow and 
elongate, e.xtending as a furrow to the anterior margin of the head. 

Key to Genera of the Suborder Amblycera. 

A. Tarsi vrith one claw; infesting mammals only (family Gyropidce) Gyropus. 

AA. Tarsi with two claws; infesting birds only (family Liotheidce). 
B. Ocular emargination distinct, more or less deep. 

Book-lice and Bark-lice; Biting Bird-lice 119 

C. Forehead evenly rounded, without lateral swellings; antenna; projecting 

slightly beyond border of the head Colpocephalum. 

CC. Forehead with strong lateral swellings. 

D. Mesothorax separated from metathorax by a suture. .. .Trinoton. 

DD. Meso- and metathorax fused; no suture L^mobothrium. 

BB. Ocular emargination absent or very slight. 

C. Sides of the head straight or slightly concave, with two small laterally 

projecting labral lobes Physostomum. 

CC. Sides of the head sinuous; forehead without labral lobes. 

D. Ocular emargination filled by a strong swelling; sternal markings 

forming a quadrilateral without median blotches Nitzschia. 

DD. Ocular emargination without swelling, hardly apparent or entirely 
lacking; median blotches on sternum. 

E. Very large; with two-pointed appendages on ventral aspect 
of hind head; anterior coxje with very long lobe-like append- 
ages Ancistrona. 

EE. Small or medium; without bi-partite appendages of hind head. 


The Mallophaga most likely to come under the observation of people 
not collectors of birds are the species which infest domestic fowls and mam- 
mals, and the following few descriptions and figures of particular species 
are therefore limited to such kinds. 

The most notorious member of the order is the common chicken-louse, 
Menopon pallidum (Fig. 146). It is of a pale straw-yellow color, from 
I mm. (^ig- in.) to 1.5 mm. 
in length, and is an un- 
usually swift and active 
little pest. Other Mallo- 
phaga infesting chickens 
are Goniocotes hologaster, 
recognized by its squarish 
head with angulated 
temples, and Lipeiirus 
variabilis, 2 mm. (y^g- in.) 
long and slender, with dis- 
tinct black markings on 
the otherwise smooth, 
white body. 

Ducks are infested by 
several species. Com- ^^^ ^^^ p^^ ^^^ 

mon among them is the Fig. 146.— The biting chicken-louse, Menopon pallidum. 
little Docophoriis icterodes _ (After Piaget; natural size, i to 1.5 mm.) 

(Fig. 147), I mm. (uV i"-) 
long, with head curiously 
expanded and rounded in front, darkish-red head, and thorax with darker 

icterodes. (Natural size indicated by line.) 

I20 Book-lice and Bark-lice; Biting Bird-lice 

bands, and a white region in the middle of the abdomen. Trinoton luridiim 
is another common duck-louse unusually large, being from 4 to 5 mm. (5 in.) 
long and readily distinguished by the triangular 
head with lateral swellings, and the abdomen with 
pronounced blackish-brown transverse bands. 

Fig. ISO. 


Fig. 152. 

Fig. 148. — A biting louse of pigeons, Lipeurus bacillus. (Natural size indicated by line.) 

Fig. 149. — Biting louse of the dog, Trichodedes latus. (After Nitzsch; natural size, 
I to 1.5 mm.) 

Fig. 150. — Biting louse of the horse, Trichodcctes parumpilosus, male. (After Morse; 
natural size shown by line.) 

Fig. 151. — Biting louse of cattle, Trichodcctes scalaris. (After Lugger; natural size, 1.5 
to 2 mm.) 

Fig. 152. — Biting louse of fringilline birds, Docophorus communis. (Natural size in- 
dicated by line.) 

Book-lice and Bark-lice; Biting Bird-lice 121 

Pigeons are almost always infested by a long and very slender louse, 
Lipeuriis baculus (Fig. 148). The head and thorax are reddish brown, 
while the abdomen is dusky with darker segmental blotches. This bird- 
louse was described and named more than two hundred years ago. 

All of the species infesting domestic mammals belong to the genus Tricho- 
dectes. Dogs are often infested by Trichodectes latus (Fig. 149), a short, 
wide-bodied species about i mm. long; while cats are less often infested by 
T. subrostratus, distinguishable by the rather pointed head with a short, 
longitudinal furrow on the under side. Horses and ^4flonkeys are troubled 
by two or three species, of which T. pilosiis, a hairy form with antenna? rising 
near the front of the head, and T. pariim pilosiis (Fig. 150), a broader-bodied 
form with head larger and less flatly rounded in front, are most common. 
Trichodectes scalaris (Fig. 151) infests cattle the world over, while sheep 
and goats have species peculiar to themselves. Comparatively few species 
of Trichodectes have been recorded 
from wild mammals, but this is 
simply because they have not been 
sought with care. Species have 

153- Fig. 154. 

Fig. 153. — A biting louse of gulls, Nirmus felix, male. (Natural size indicated by line.) 
Fig. 154. — Giant bird-louse of the albatrosses, Ancistrona gigas, male. (Natural size 
indicated by line.) 

been found on the bear, raccoon, fox, coyote, weasel, gopher, beaver, deer, 
skunk, and porcupine. Gyropus, the other mammal-infesting genus of 

122 Book-lice and Bark-lice; Biting Bird-lice 

Mallophaga, has been found only on the guinea-pig. Washing the body of 
the infested animal with kerosene emulsion (see p. 190) is probably the 
most effective remedy for biting lice. 

Of the nearly three hundred species of Mallophaga which I have recorded 
(Proc. Nat. Mus., v. 22, 1899, pp. 39-100) from wild North American birds, 
mention may be made of the largest, Lcemobothnum loomis, taken from the 
Canada goose; of Docophorus communis (Fig. 152), the most abundant and 
widely distributed parasite of perching and song birds; of the pretty Nirmns 
jelix (Fig. 153), with its clean white body and sharply marked black spots; 
of the fierce-looking Lipeurus ferox, found on albatrosses; and of Ancistrona 
gigas (Fig. 154), found on fulmars, the broadest of the Mallophaga. 

As there are nearly one thousand different species of North American 
birds, and Mallophaga have been taken from but two hundred and fifty of 
them, it is obvious that the collector and student of these parasites has a 
profitable field open to him. 



(Order Orthoptera) 

do not shut up our singing insects in cages 
as the Japanese do, and bring them into 
the house to cheer or amuse us, but we do 
enjoy them, and were our summer and 
early fall days and nights to become sud- 
denly silent of chirping and shrilling, we 
should realize keenly how companionable 
crickets and grasshoppers and katydids 
had been for us. A wholesome blitheness 
and vigor and ecstasy of living rings out 
in the swift and steadfast song of most of 
our field and wood insect singers, while 
the cheeriness of the cricket on the hearth 
is familiar poetry and proverb. 

Almost all this insect music comes from the members of one order, the 
Orthoptera. Indeed there is but one famous insect maestro, the cicada (of the 
order Hemiptera), which does not belong to the group of crickets, locusts, green 
grasshoppers, and katydids. Besides being singers, too, the Orthoptera 
are the characteristic leapers of the insect world; crickets and locusts easily 
surpass the world's athletes for high jumping if the record takes into account 
the comparative size of the athletes. And, curiously, the singing Orthoptera 
are the leaping ones. Of the six families composing the order, three include 
insects which do not sing nor leap, while the other three are made up of 
singers and leapers. 

As one tramps the roadways or dry pastures in summer and autumn, 
the steady shriUing of the locusts on the ground, or their sharp "clacking" 
as they spring into air, are most familiar sounds. When you ramble through 
the uncut meadows and lush low grounds the still shriller singing of the 
slender-bodied, thin-legged, meadow green grasshopper is heard, while 
in the orchards and woods the snowy tree-crickets and broad-winged katydids 


I 24 Cockroaches, Locusts, Grasshoppers, and Crickets 

keep up 
offers its 

the chorus. At home, in house and garden, the domestic cricket 
music to the already over-full ears. All this choiring is done by 
singers without a voice; that is, without the 
production of sound from the throat and 
mouth by means of vocal cords set into vi- 
bration by air. Insects are orchestral per- 
formers, using their legs and wings, for the 
most part, to make their music. When the lo- 
cust sings while at rest, it is rasping the inner 
surface of the broad hind thighs across the 
roughened outer surface of the folded fore 
wings; when it "clacks" in the air, it is strik- 
ing the front margin of the hind wing back 
and forth past the hinder margin of the 
thickened fore wings. When the cricket 
shrills on the hearth, or anywhere else, he, for 
only the male crickets have the musical gift, is holding 
his fore wings up over his body at an angle with it of 
about 45° and is rubbing together the upper surfaces of 
the basal region of the fore wings, which are specially 
modified for this purpose. The tree-crickets, katydids, 
and meadow green grasshoppers have, in the males, 
the same general sort of music-making apparatus as 
the cricket, and sing by similarly rasping or rubbing 
together the modified parts of the fore wings. This 

Fig. 155. — Longitudinal section through head and neck of locust, 
showing disposition of alimentary canal, brain, and sub- 
cesophageal ganglion. (After Snodgrass; much enlarged.) 

music-making by rasping is called stridulation, and for the most part 
insect stridulation is strictly strident, and sounds to better advantage in the 
field than it would from caged songsters in the parlor. 

Cockroaches, Locusts, Grasshoppers, and Crickets 125 

All the Orthoptera have biting mouth-parts, and bite off and chew their 
food, which is usually live vegetable matter, especially green leaves, 
although the members of one family are predaceous, preying on other insects, 
and those of another family prefer dried vegetable or animal matter. The 
metamorphosis is incomplete, the young, when hatched, resembling the parents 
except for small size and lack of wings. The young have the same feeding 
habits and same haunts as the adults, and by development and growth the 

Fig. 156. — The immature stages of a locust, Melanoplus jemur-rubrum. a, just hatched, 
without wing-pads; h, c, d, and e after first, second, third, and fourth rnoultings 
respectively, showing appearance and development of wings; /, adult, with fully 
developed wings. (After Emerton.) 

wings and parental stature are soon acquired. The name of the order is 
derived from the straight-margined leathery fore wings, or elytra, whose 
chief function is to cover and protect the larger membranous hind wings 
on which the flight function depends. Among the leaping Orthoptera the 
hindmost legs are very large and long, and when at rest or in walking the 
"knee-joints" of these legs are much higher than the back of the insect. 

The three singing and leaping families are the AcridiidEe, locusts and 
grasshoppers with short antenna;; Locustidae, meadow green grasshoppers 

126 Cockroaches, Locusts, Grasshoppers, and Crickets 

and katydids, all with long thread-like antenna?; and Gryllidae, the crickets. 
The three silent and walking or running families are the Blattidae, cock- 
roaches; Mantidae, praying-horses and soothsayers; and Phasmidas, walk- 
ing-sticks or twig-insects. These families can be distinguished by the follow- 
ing table: 


Non-leaping and mute; hind femora closely resembling those of the other legs and 
scarcely stouter or longer than the middle femora; tarsi 5-segmented; ovipositor 
Body oval, depressed; head nearly horizontal and nearly or quite concealed by 

the flattish shield-like pronotum; quickly running (Cockroaches.) Blattid.e. 

Body elongate, generally narrow; head free, often with constricted neck; pronotum 
elongate, never transverse; slowly walking. 
Fore legs spined and fitted and held for grasping; antennae usually shorter than 
body; pronotum usually longer than any other body segment; anal cerci 

jointed (Praying Mantes.) Mantid^. 

Fore legs not fitted for grasping; antennae usually longer than body; pronotum 

short (Leaf -insects and Walking-sticks.) Phasmid.i:. 

Leaping and usually capable of stridulation; hind femora stouter or longer, or both, 
than the other femora; the hind legs enlarged, for leaping; tarsi 4- or 3-segmented; 
head vertical; ovipositor usually visible. 
Antenna; much shorter than the body (with few exceptions); ocelli three; tarsi 3-seg- 
mented; auditory organs, when present, situated on basal abdominal segment; 
ovipositor composed of two pairs of short, strong, slightly curving pieces. 

(Locusts.) ACRIDIIDvE. 

Antennaj much longer than the body, delicately tapering; tarsi 3- or 4-segmented; 
auditory organs usually near the base of the fore tibise; ovipositor usually pro- 
longed into a compressed blade, or needle, its parts compact. 
Tarsi 4-segmented; ocelli usually absent; ovipositor usually exserted and forming 
a strongly compressed, usually curving, blade with tip not expanded. 

(The long-horned grasshoppers.) Locustid^. 
Tarsi 3-segmented; ocelli variable; ovipositor usually exserted and forming a 
nearly cylindrical straight needle, the tip somewhat expanded. 

(Crickets.) Gryllid.e. 

Mrs. Smith takes it amiss when you ask permission to collect "roaches" 
in her house, and will prove to you any day the conspicuous absence of these 
unwelcome guests in the scrubbed and spotless pantry and kitchen. But 
with a candle go stocking-footed at night into the same kitchen and you 
will not unlikely find "good hunting." Although but few of the thousand 
different kinds of cockroaches known in the world are to be found in the 
United States, these few, and particularly three or four imported foreigners 
among them, are very abundant, and, after dark, very much in evidence in 
their favorite habitat. Their chosen abiding-place is in kitchens, pantries, 
laundries, restaurants, bakeshops, etc., where the atmosphere is warm 

Cockroaches, Locusts, Grasshoppers, and Crickets 127 

and humid and the roach's table is well set with good things. Almost any 
sort of dry organic matter suits their taste; bread, crackers, miscellaneous 
cold-lunch delicacies, the paste of bookbindings and wall-paper, leather, 
woolens, and even their own egg-cases and cast skins making up the dietary. 
The fo'k'sel and galley of ships are the roaches' special joy; the hotels and 
restaurants of tropic and subtropic lands house swarms of these bill-evadint^ 
guests. From Mazatlan, Mexico, a naturalist sent me quarts of large native 
American roaches (Periplaneki americana), which he readily scooped up 
from his bedroom floor. Ships come into San Francisco from their long 
half-year voyages around the Horn with the sailors wearing gloves on their 
hands when asleep in their bunks in a desperate effort to save their finger- 
nails from being gnawed off by the hordes of roaches which infest the 
whole ship. A few of our species still live outside under stones and old 
logs, but most of them have learned that an easier life awaits them in the 

The roaches compose the Orthopterous family Blattida^, and are an 
ancient and persistent insect group. In Carboniferous times, before flies, 
butterflies, bees, and wasps had come into existence, cockroaches were 
the dominant insects. The body in all is flattened and slippery with the 
legs adapted for quick running, so that the insects are well fitted to escape 
safely into narrow cracks and crevices. The head is concealed from above 
by the expanded shield-shaped dorsal wall of the prothorax (pronotum). 
Wings are present in most species, the front pair 
leathery and serving, when the wings are folded, to 
cover and protect the larger, thin, membranous 
hind pair. In some forms the females are wingless, 
and the indoor habit may be held responsible for 

the lessened usefulness and resultant loss of the ^^^- ^S7- — £gg-case of 
'-T'l- ii- i. c^^ J r i-'i.- 1 1 cockroach. (Three times 

wmgs. 1 he mouth-parts are fitted tor bitmg hard natural size ) 

dry substances, the jaws being strong and toothed. 

The eggs are laid in small purse-like, horny, brown cases (Fig. 157), which 

are usually carried about by the female until the young are ready to issue. 

The young grow slowly, requiring probably about a year, in most species, 

to become fully developed. From the beginning, the young can run about 

and take care of themselves, eating the same kind of food as the adults. 

They moult several times during growth, and at each moult the wing-pads 

are a little larger. 

There are four common species of cockroaches found in dwellings in this 

country, only one of which is native. This is the large American roach, 

Periplaneta americana, about i| inches long (to tip of folded wings), light 

brown in color, and with the wings expanding nearly 3 inches. This species 

is abundant in the middle and western states, having gradually extended 

128 Cockroaches, Locusts, Grasshoppers, and Crickets 

its range north from its native region in Mexico and Central America. The 
Australian roach, Periplaneta australasia, resembles P. americana, but is 
darker in ground color, a qusrter of an inch shorter, and has a conspicuous 
yellow submarginal band running around the shield-shaped pronotum. 
Each fore wing has also a strong yellow tapering bar in the basal part of 
the costal region. It came originally from the Australian Pacific region, 
and is now spread widely over the world, being common in this country 
in Florida and other southern states. The most abundant and destruc- 
tive house-roach in the eastern states is the small German cockroach, 
Ectohia gennanica (Fig. 158), about half an inch long, and pale yellowish 
brownish with a pair of distinct black longitudinal stripes on the pro- 

Fig. 158. Fig. 159. Fig. 160. 

Fig. 158. — The croton-bug, or German cockroach, Ectohia gennanica. (Twice natural 

Fig. 159. — The black beetle, or Oriental cockroach, Periplaneta orientalis. (One and 

one-half times natural size.) 
Fig. 160. — The common wood cockroach, Ischnoptera pennsylvanica. (After Lugger; 

natural size indicated by line.) 

notum. This roach is often called croton-bug, from its intimate asso- 
ciation with the pipes of New York City's Croton-water system. It is an 
importation from Europe, where it is especially abundant in Germany. Its 
real nativity is unknown, but it is now of world-wide di.stribution. The 
fourth species is the black or Asiatic roach, or black beetle, as it is sometimes 
called, Periplaneta orientalis (Fig. 159). This roach is about one inch 
long, with brownish-black body; in the female the wings are rudimentary, 
and in the male the wings when folded do not quite reach the tip of the 
abdomen. This species is common in all the eastern and Mississippi 
Valley states and extends as far west as the great plains. It is the 
commonest cockroach in England and Europe. The native outdoor species 
most familiar in this country is the common wood-cockroach, Ischnoptera 

Cockroaches, Locusts, Grasshoppers, and Crickets 129 

pennsylvanica (Fig. 160), with long, light-colored wing-covers, and wings 
which extend considerably beyond the tips of the abdomen. The margin 
of the pronotum is light, the disk being dark, and the front margins 
(lateral when folded) of the wing-covers are lighter than the discal 
parts. The body is an inch long and rather narrow and slender. This 
species is common in the woods and sometimes comes into houses in 

In the southern states and those of the Mississippi Valley a large insect 
may be not infrequently seen standing motionless in a corner of a window, 
in a striking attitude. This attitude may be taken as one of hopeful prayer, 
as those who gave the name praying-mantis to the insect seem to have taken 
it, or one of self-confident readiness to do violent work with those upraised, 
sharply spined, and very willing fore feet. This is the way the house-flies 
rightfully take the mantis's attitude. Watch an unwary bluebottle crawl or 
buzz into the fatal corner. Blundering buzziness is finished for that blue- 

FiG. 161. — The praying-mantis, Mantis religiosa. (After Slingerland; natural size.) 

bottle; and the first course of a square meal has come to him who waits 
and watches. Other names, as rearhorse, camel-cricket, and soothsayer, 
have been given the mantis, all suggested by the attitude and curious body 
make-up of the creature. The prothorax is long and stem-like, the head 
broader than long, with protuberant eyes, and the fore legs are not used 
for locomotion, but are large, strongly spined, and fitted for seizing and hold- 
ing the prey. The wings are short and broad and usually rather leaf-like 
in coloration and texture, the whole insect when at rest resembling somewhat 
a part of the plant on which the mantis ordinarily stands. The window-corner 
is a new and unnatural locale for the insects, but the abundance of prey here 
in summer-time makes it a good feeding-ground. 

The family Mantidc-e includes less than a score of species in this country, 
all of them southern in range, and only a few occurring north of the Rio 

130 Cockroaches, Locusts, Grasshoppers, and Crickets 

Grande and Gulf coast regions. All the species are carnivorous, and 

undoubtedly do much good in making away with many noxious insects. In 

1899 some specimens of the common European praying-mantis, Mantis 

religiosa (Fig. 161), were found in and 
near Rochester, N. Y. They had 
probably been accidentally imported 
into this country in nursery stocks from 
France. As this species seems able 
to live farther north than our native 
species, Professor Slingerland is laud- 
ably trying to establish it in our coun- 
try. He takes care of a colony, and 
is distributing many of the egg-cases 
over the entire country. All the man- 
tids lay their eggs in curious masses 
(Figs. 162 and 163), covered with a 
quickly drying tough mucus. These 
egg-cases are attached to branches and 
plant-stems in the fall, and the young 
hatch in the following summer and 
soon grow (moulting several times 
and developing wings) to full stature, 
which for our most common native 
species, Stagmomantis Carolina, is 
about 2^ inches long. 

Slingerland has collected a num- 
ber of the old accounts of the Euro- 
pean mantis which are of interest as proofs of the light and graceful fancy 

of some of the early author-naturalists. 

The ancient Greeks gave the insects 

the name Mantis, that is, ** prophet." 

Mouffet, writing over five hundred 

years ago, says: "They are called 

Mantes, that is, Joretellers, either 

because by their coming (for they first 

of all appear) they do show the spring 

to be at hand, so Anacreon the poet 

sang; or else they foretell death and Fig. 163. — Egg-case of praving-mantis, 

famine, as Cslius the Scoliast of ^^'""'' ''^TT' "'"' T""' rST'"f 
' ■^'^ ^^ arrangement of eggs inside. (Natural 

Theocritus has observed ; or, lastly, size ) 

because it always holds up its fore feet like hands praying as it were, after 

the manner of their Diviners, who in that gesture did pour out their sup- 

FiG. 162. — Egg-cases of the praying- 
m-antis, Mantis religiosa. (After 
Slingerland; natural size.) 

Cockroaches, Locusts, Grasshoppers, and Crickets i 3 1 

plications to their Gods." And he says again: "They resemble the Diviners 
in the elevation of their hands, so also in likeness of motion; for they do not 
sport themselves as others do, nor leap, nor play, but walking softly, they 
retain their modesty, and shewes forth a kind of mature gravity. ... So 
divine a creature is this esteemed, that if a childe aske the way to such a 
place, she will stretch out one of her feet, and shew him the right way, and 
seldome or never misse." Piso in his works states that mantids "change into 
a green and tender plant, which is of two 
hands' breadth. The feet are fixed into 
the ground first; from these, when neces- 
sary, humidity is attracted, roots grow out 
and strike into the ground; thus they 
change by degrees, and in a short time 
become a perfect plant." 

Almost everywhere that mantids occur, 
strange superstitions are held concerning 
them. Most of these ascribe some degree 
of sanctity to them, and to kill them 
maliciously is considered sinful. Cowan 
says that "the Turks and other Moslems 
have been much impressed by the actions 
of the common Mantis religiosa, which 
greatly resemble some of their own attitudes 
of prayer. They readily recognize intelli- 
gence and pious intentions in its actions, 
and accordingly treat it with respect and 
attention, not indeed as in itself an object 
of reverence or superstition, but as a fel- 
low worshipper of God, whom they believe 
that all creatures praise with more or less 
consciousness and intelligence. Other su- 
perstitions with respect to the Mantis are 
current: when it kneels it sees an angel 
in the way, or hears the rustle of its wings; 
when it alights on your hand you are about 
to make the acquaintance of a distin- 
guished person; if it alights on your head, 

a great honor will shortly be conferred Fig. 164. — The walking-stick, Diaphe- 

H-. ■ ■ ■ romera jemorata. 

it mjures you ni any way, 

which it does but seldom, you will lose a valued friend by calumny. Never 

kill a Mantis, as it bears charm against evil." Finally, monkish legends 

tell us, says Slingerland, that St. Francis Xavier, seeing a Mantis moving 

132 Cockroaches, Locusts, Grasshoppers, and Crickets 

along in its solemn way, holding up its two fore legs as in the act of devo- 
tion, desired it to sing the praise of God, whereupon the insect carolled 
forth a fine canticle! 

More amazing than the Mantids for modification of form and appear- 
ance away from the usual insect type are the members of the family Phas- 
midse. The only representatives of this family in the United States are 
the walking-sticks, or twig insects (Fig. 164), of which half a dozen genera, 
with from one to three species each, have been recorded. The only one 
of these genera which is found in the East is Diapheromera, of which D. 
jemorata is the common species. Our other Phasmids are found in the 
West or extreme South. All of our species are wingless and are generally 
sluggish in movement, and depend for protection largely on their amazingly 
faithful resemblance in shape and color to twigs, and on their capacity to 
emit an ill-smelling fluid from certain glands on their prothorax. Diaphero- 
mera jemorata (Fig. 164) feeds on the leaves of oaks, walnuts, and probably 
other trees. It drops its hundred seed-like eggs loosely and singly on the 
ground, where they lie through the winter, hatching irregularly through 
the following summer. Some may even go over a second winter before 
hatching. Fernorata may be either brown or green; so it frequents dead 
or leafless, or live and green-leaved parts, according to the correspondence 
of its body color with the one or the other of these environments. The long, 
slender, wingless body, the thin, long legs held angularly, and the harmonizing 
body color, all serve to make the walking-stick well-nigh indistinguishable 
when at rest on the twigs. 

In tropic and subtropic countries the Phasmids are numerous (over 600 
species are known) and present other striking resemblances to the details 
of their habitual environment. A conspicuous and perfect example of 
resemblance is the green leaf-insect Phyllium (PI. XIII, Fig. 2), whose wings, 
flattened body, and expanded plate-like legs, head, and prothorax, all bright 
green and flecked irregularly with small yellowish spots, like those made 
by the attacks of fungi on live leaves, combine to simulate with wonderful 
effect a green leaf. 

Other examples of such protective resemblance and a discussion of the 
origin and significance of the phenomenon may be found in Chapter XVII 
of this book. 

The genera of Phasmidae occurring in the United States may be distin- 
guished by the following key: 

Tibiae with a groove at tip to receive the base of the tarsi when bent upon them. 
Antennae with less than twenty segments, and much shorter than the fore femora. 


Cockroaches, Locusts, Grasshoppers, and Crickets 133 

Antcnnre with many segments, and longer than the fore femora. 

Mcsothorax twice as long as prothorax Anisomorpha. 

Mesothorax no longer than prothorax Tinema. 

Tibias without groove at tip, as above described. 

Hind femora with one or more distinct spines on the median line of the under side 

near the tip Di apheromera. 

Hind femora without such spines. 

Head, especially in female, with a pair of tubercles or ridges on the front between 

the eyes Sermyle. 

Head without such tubercle or ridges Bacunculus. 


One day in early summer of the Centennial Year (1876) the people all 
over Kansas might have been seen staring hard with shaded eyes and serious 
faces up towards the sun. By persistent looking one could see high in the 
air a thin silvery white shifting cloud or haze of which old residents sadly 
said, "It's them again, all right." Now this meant, if it were true, that, 
far from being all right, it was about as wrong as it could be for Kansas. 
"Them" meant the hateful Rocky Mountain locusts, and the locusts meant 
devastation and ruin for Kansas crops and farmers. In 1866 and again 
in 1874 and 1875 the locusts had come; first a thin silvery cloud high over- 
head — sunlight glancing from millions of thin membranous fluttering 
wings — and then a swarming, crawling, leaping, and ever and always 
busily eating horde of locusts over all the green things of the land. And 
the old residents spoke the truth in that summer of 1876. It was "them," 
uncounted hosts of them, and only such patriotic farmers as had laid by 
money for a rainy day or a grasshopper year could visit the Centennial 

Not all locusts are migratory or appear in such countless swarms as 
this invader from the high plateau of the northern Rocky Mountains. In 
South America another locust species, larger than ours, has similar habits; 
having its permanent breeding-grounds on the great plateau at the eastern 
foot of the Chilean Andes and descending almost every year in swarms on 
the great wheat-fields of Argentina. And in Algeria and Asia Minor occurs 
the migratory locust of the Scriptures, a still other and larger species. But 
of the 500 (app.) locust species, members of the family Acridiidae, which 
are known in the United States but three or four can be fairly called 
migratory, and of these the Rocky Mountain locust, Melanopliis sprelus, is 
the most conspicuous. The lesser migratory locust, Melanopliis atlanis, 
does much injury in New England and other eastern states, while the 
pellucid locust, Camnula pellncida, is a migratory species that often does 
much harm in CaHfornia and other western states. Sometimes large 
bodies of immature wingless individuals of the large species Dissosteira 
longipennis, abundant on the plains of eastern Colorado and western Kansas 

134 Cockroaches, Locusts, Grasshoppers, and Crickets 

will move slowly on, walking and hopping for many miles, eating every 

green weed and grass-blade in their path, but this is only a limited and 

local sort of migration. 

Almost all the Acridiidae, despite the many species in the family, are 

readily recognizable as locusts 

or grasshoppers — short-horned 

grasshoppers they may be called, 

to distinguish them from the 

meadow green grasshoppers with 

long thread-like antennae — because 

of their general similarity in ap- 

„ . T w 1.1 WW. • pearance and habit. The body 

Fig. 105. — Locust from lateral aspect (left wings f •' 

removed), showing (ao.) auditory organ, is rather robust, the head is set 
(Natural size.) ^,^1-]^ j^g joj^g g^^is at right angles 

with the axis of the body, so that the mouth with its strong biting and 
crushing jaws is directed downwards (Fig. 165); the antennae are never 
as long as the body and are composed of not more than twenty-five 
segments; the prothorax is covered laterally as well as dorsally by its large 
saddle-like horny pronotum, which projects so as also to cover and protect 
from the sharp grass-blades the soft thin-walled neck and the equally 
thin-walled suture between prothorax and mesothorax; the abdomen is 
broadly and closely joined to the metathorax, and 
in the female ends in a short and strong ovipositor 
composed of four horny pointed pieces; the hind 
legs are much larger than the others and fitted 
for leaping, and the fore wings, called tegmina, 
are narrow and straight-margined, and serve 
specially to cover and protect the much larger 
thin membranous hind wings, which are plaited 
and folded like a fan when the locust is at rest. 

The sounds or stridulation of locusts are 
made in two ways. When at rest certain species 
draw the hind legs up and down across the wing- 
covers so that numerous fine little ridges on the 

inner surface of the broad femora are rasped pj^^g^^Locust impaled on 
across a thickened and ridged longitudinal vein thorn by shrike (butcher- 
on the outer surface of the wing-covers. When ^^''^^- (Natural size.) 
in flight certain locusts rub or strike together the upper surface of the 
front edge of the hind wings and the under surface of the fore wings 
or tegmina. This produces a loud, sharp clacking which can be heard 
for a distance of several rods. The loudest "clacking" of this kind 

Cockroaches, Locusts, Grasshoppers, and Crickets 135 

that I have heard is made by a species of Trimerotropis, abundant in 
the beautiful little glacial "parks" of the Colorado Rockies. Locusts 
undoubtedly make sounds to be heard by each other, and it is not difficult 
to find in them — a matter of more difficulty in most other insects — certain 
organs which are almost certainly auditory organs, or ears. On the outer 
faces of the upper part of the first abdominal segment is a pair of sub- 

FiG. 167. — The red-legged locust, Melanoplus jemiir-rubrum, female. 
(After Lugger; natural size indicated by line.) 

circular clear window-like spots (Figs. 165 and 55). These are thin places 
in the body- wall serving as tympana; on the inner face of each is a small 
vesicle, and from it a tiny nerve runs to a small auditory ganglion (nerve- 
center) at one side of the tympanum. From this auditory ganglion a nerve 
runs to the large ventral ganglion in the third thoracic segment. Similar 
auditory organs are found in the other singing Orthoptera, the crickets and 
katydids, but situated in the front legs instead of on the back. 

136 Cockroaches, Locusts, Grasshoppers, and Crickets 

The life-history of all our locusts is, in general characteristics, very similar. 
The eggs are deposited in oval or bean-shaped packets enclosed in a glutin- 
ous substance. They are usually laid just below the surface of the soil, 
but in some cases are simply pushed to the ground among the stems of 
grasses, while a few locust-species thrust them into soft wood. The strong, 
horny ovipositor at the tip of the abdomen is worked into the ground, the 
four pieces separated, and the eggs and covering mucous material extruded. 
The eggs in a single mass number from twenty-five to one hundred and 
twenty-five, varying with different species, and the females of some species 
lay several masses. The different species also select different times and 
places for egg-laying, some ovipositing in the fall and some in the spring, 
while some select hard, gravelly, or sandy spots or well-traveled roads, and 
others choose pastures and meadows and the uncultivated margins of irriga- 

If the eggs are laid in the fall, the more usual case, they do not hatch 
until the following spring. The young hoppers are of course wingless, very 
small, and pale-colored, but they have the general body make-up of their 
parents, with the biting mouth and long-leaping hind legs. They push 
their way above ground and feed, as do the adults, on the green foliage of 
grasses, herbs, or trees, and in two or three months become full grown and 
mature, having moulted five or six times during this growth and developed 
wings. The wings begin to appear as minute scale-like projections from 
the posterior margins of the back of the meso- and meta-thoracic segments, 
and with each moulting are notably larger and more wing-like in appear- 
ance. During all this development the wing-pads are so rotated that the 
hinder wings (always underneath the fore wings in the adult locust) lie out- 
side of and above the fore wings (Fig. 156). 

The family Acridiidae includes in the United States about 500 species, 
representing 107 genera. These genera are grouped in four subfamilies 
as follows: 


Pronotum (dorsal wall of prothorax) extending back over the abdomen nearly or quite 

to its tip; tegmina (fore wings) short and scale-like Tettigin^. 

Pronotum not extending back over abdomen or only slightly; tegmina usually well 
developed (sometimes short or wanting). 
Prosternum (ventral aspect of prothorax) with a prominent thick conical or cylindrical 

spine AcRiDiiNiE. 

Prosternum not spined (sometimes a short, oblique, inconspicuous, obtuse tubercle). 

Face very oblique Tryxalin^. 

Face nearly or quite vertical (Edipodin^. 

In the subfamily Acridiinae the most conspicuous and economically 
important member is the Rocky Mountain or hateful migratory locust, 

Cockroaches, Locusts, Grasshoppers, and Crickets 137 

Melanopliis spretiis. The invasions of the grain-growing Mississippi Valley 
states by this species have been already mentioned. In 1866, 1874, and 
1876 such invasions occurred, and before these still others. "Kansas grass- 
hoppers" had gained a notoriety which spelled ruin to the state. But, 
strangely, these grasshoppers, or locusts, not only were not Kansas born, 
but could not even adopt Kansas as a home. The Rocky Mountain locust 

Fig. 168. Fig. 169. 

Fig. 168. — The lesser migratory locust, Melanopliis atlanis, female. (After Lugger; 

natural size indicated by line.) 
Fig. 169. — The differential locust, Melaiioplus differentialis, female. (After Lugger; 

natural size indicated by line.) 

has its permanent breeding-grounds on the plains and plateaus of Colorado, 
Idaho, Wyoming, Montana, and British Columbia, at an altitude of from 
2000 to 10,000 feet above sea-level, and while able to maintain itself for 
a generation or two in the low, moist Mississippi Valley, cannot take up 
any permanent residence there. But in those days there were few ranches 
and farms on the great plains, and succulent corn and wheat were not at 

138 Cockroaches, Locusts, Grasshoppers, and Crickets 

hand to feed the milUons of young which hatched each spring. So, after 
exhausting the scanty wild herbage of their breeding-grounds, and develop- 
ing to their winged stage, hosts of locusts would rise high into the air until 
they were caught by the great wind-streams bearing southeast, and, with 
parachute-like wings expanded and air-sacs in the body stretched to their 
fullest, would be borne for a thousand miles to the rich grain-fields of the 


170. — The two-striped locust, Melanophis hivittatus, female. 
(After Lugger; natural size indicated by line.) 

Mississippi Valley. As far east as the middle of Iowa and Missouri and 
south to Texas these great swarms would spread; and once settled to ground 
and started at their chief business, that of eating, not a green thing escaped. 
First the grains and grasses; then the vegetables and bushes; then the 
leaves and fresh twigs and bark of trees! A steady munching was audible 
over the doomed land! And this munching was the devouring of dollars. 
Fifty millions of dollars were eaten in the seasons of 1874-76 alone. 

Cockroaches, Locusts, Grasshoppers, and Crickets 139 

Remedies there were practically none; when the summer hosts laid 
their eggs in the ground for the one generation that could be reared in the 
invaded land, these eggs could be plowed up, a remedy that is used with 
much success in the far western locust-infested states; also when the wingless 
voung "hoppers" appeared in the spring they could be crushed by heavy 



-The American locust, Schistocerca americana, female. 
(After Lugger; natural size.) 

rollers drawn across the fields by horses, or burned by scattering straw over 
the helpless host and lighting it. Both of these remedies are also used in 
western locust-fighting. But against the winged adults there is little that 
can be done. 

In Asia and South America, where there are also migratory locusts (of 
different, much larger species) the natives sometimes try to frighten away 
an alighting swarm by smoke and noise, but such a swarm as that which 
passed over the Red Sea in November, 1889, spread out for over 2000 square 

140 Cockroaches, Locusts, Grasshoppers, and Crickets 

miles in area, would be little affected by a bonfire. In Cyprus in 1881, 
1300 tons of locust-eggs were destroyed; how many eggs go to make a ton 
one can only faintly conceive of. 

There has been no serious Rocky Mountain locust invasion of the Missis- 
sippi Valley since 1876, and there will probably never be another. The 
locust is being both fed and fought in its own breeding range; many are 

Fig. 172. Fig. 175. 

Fig. 172. — The emarginate locust, Schistocerca emarginata, male. (After Lugger; nat- 
ural size.) 

Fig. 173. — The pale-green locust, Hesperotettix pratensis, female. (After Lugger; 
natural size indicated by line.) 

Fig. 174. — The short-winged locust, Stenobothrus curtipennis, female. (After Lugger; 
natural size indicated by line.) 

Fig. 175. — The sprinkled locust, Chloealtis conspersa, male. (After Lugger; natural size 
indicated by line.) 

killed every year, and for those that are left there is food enough and to spare 
in the great grain-fields of the northwest plains. 

The genus Melanoplus, to which the Rocky Mountain locust belongs, 
is the largest of all our Acridiid genera, one hundred and twenty species 
found in the United States belonging to it. Of these species a very common 
one all over the country is the red-legged locust, Melanoplus jemur-riihrum 
(Fig. 167), which is about one inch long, with olivaceous brownish body, 
clear hind wings and brownish fore wings that have an inconspicuous 
longitudinal median series of black spots in the basal half (these spots 

Cockroaches, Locusts, Grasshoppers, and Crickets 141 

sometimes wanting). The hind tibiae are normally red (sometimes yellow- 
ish), hence the name, although these red hind legs are common to many 
other locust species. The lesser migratory locust, M. allanis (Fig. 168), 
is a species of about the same size and appearance which sometimes 
appears in great swarms and does much injury to crops. The largest 
species of the genus is M. difjerentialis (Fig. 169), over an inch and a half long, 
with brownish-yellow body, fore wings without spots, and hind wings clear. 
It is common in the Southwest, where, in company with M. bknttatus (Fig. 
170), nearly as large but readily distinguished by the pair of longitudinal 

Fig. 176. Fig. 177. Fig. 178. 

Fig. 176. — The short-winged green locust, Dichromorplia viridis, female. (After Lugger; 

natural size indicated by line.) 
Fig. 177. — The spotted-winged locust, Orphida pelidina. (After Lugger; natural size 

of male 16-19 mm., of female, 20-24 mm.) 
Fig. 178. — The Carolina locust, Dissoskira Carolina, female. (After Lugger; natural 

size indicated by line.) 

pale-yellowish stripes extending from the head across the thorax and along the 
folded wing-covers nearly to their tips, it often becomes sufficiently abundant 
to do serious injury. These two species are always to be found commonly 
in western Kansas, and bivittatiis ranges far to the north, being one 
of Minnesota's destructive species. 

Among the other genera of the subfamily Acridiinae Schistocerca is con- 
spicuous because of the large size and wide distribution of its species. The 
American locust, S. americana (Fig. 171), measures three inches from head 
to tips of tegmina, with reddish-brown body and a longitudinal yellowish 
strip extending along the head, thorax, and closed tegmina nearly to their 

142 Cockroaches, Locusts, Grasshoppers, and Crickets 

tips. The tegmina are opaque and reddish at base, subtransparent dis- 
tally; the great hind wings are clear and transparent. This locust is com- 
mon in the South, where it sometimes assumes a migratory habit and 
becomes very injurious to crops. The leather-colored locust, S. alutaceum, 
with dirty brownish-yellow body and paler stripe on top of head and thorax, 

Fig. 179. — The coral-winged locust, Hippiscus tuberculatus, female. (After Lugger; 
natural size indicated by line.) 

semi-transparent tegmina, and clear transparent hind wings, and the rusty 
locust, S. rubiginosiim, with light dust-red body and opaque tegmina, are 
the common eastern representatives of this genus. Both are large and 
striking forms. 

The subfamily Tryxalin^e includes a number of locusts distinguished 
by the sharp oblique sloping of the face, and in some cases by the much 
prolonged and pointed vertex (region of the head between the eyes). In 

the East the short-winged locust, 
Stenohothrus curtipennis (Fig. 174), 
recognizable by its short narrow 
wings, yellow under-body, and prom- 
inent yellowish hind legs with black 
knees, is a common example of this 
group. It likes to hide among tall 
grasses, where its sprightly tumbling 
Fig. 180.— Young coral-winged locust, and dodging usually save it from 
Hippiscus tuberculatus. (After Lugger; capture despite its poor flying and 
natural size indicated by line.) , . ^, 7 , , , 

leaping powers. The sprmkled 

locust, Chlivaltis conspersa (Fig. 175), is an abundant species through- 
out the East. It is light reddish brown sprinkled with black spots, 
and has pale yellowish-brown tegmina with many small dark-brown spots, 
the wings being clear; it is about three-fourths of an inch long. The 
males have the sides of the pronotum shining black. This locust lays its 
eggs in rotten stumps or other slightly decayed wood. Blatchley discovered 
a female in the act of boring a hole for her eggs in the upper edge of the 
topmost board of a six-rail fence. One of the most grotesque of all the 
locusts is a member of this subfamily named Achtinim brevipenne. The 
body is very long and thin, measuring an inch and a half in length by one- 

Cockroaches, Locusts, Grasshoppers, and Crickets 143 

tenth of an inch wide in the broadest part; the head is pointed and pro- 
jects far forward and upward, the face being very obhque. The wings 
are short and the body color brown. Comstock found this locust quite 
common in Florida on the "wire-grass" which grows in the sand among 
the saw-palmettoes, and "so closely did their brown linear bodies resemble 
dry grass that it was very difficult to perceive them." So the grotesqueness 
has its use. 

The subfamily (Edipodina^ is well represented in the United States, 

Fig. 181. — Hippiscus tigrinus, female. (After Lugger; nat. size indicated by line.) 

containing twenty- four genera and about 140 species. Almost all the familiar 
locusts with showy colored hind wings belong to this subfamily. One 
of the commonest species all over the United States and Canada is the 
Carolina locust, Dissosteira Carolina (Fig. 178), easily recognized by its 
black hind wings with broad yellow or yellowish-white margin covered with 
dusky spots at the tip. Its body color is pale yellowish or reddish brown, 
and it measures 1^-2 inches in length. It flies well; the males have the 
habit of hovering in the air a few feet above the ground and making a loud 

144 Cockroaches, Locusts, Grasshoppers, and Crickets 

Fig. 182. — The yellow-winged locust, Arpltia 
sulphiirea. (After Lugger; natural size of 
male 23-26 mm., of female 28-30 mm.) 

"clacking." The species of Hippiscus are heavy, broad-bodied forms 

with wings reddish or yellow- 
ish at base, then a broad black- 
ish band, and the apex and 
margin clear. The fore wings 
and body are yellowish to 
brown, with darker blotches 
and speckles. H. discoideus, 
with wings red on basal half, 
is common in the East. H, 
tuherculatiis (Figs. 179 and 180), 
the coral-winged locust, or king 
grasshopper, also with red 
wing-disks, is common in the 
Mississippi Valley ; it makes 
a very loud rattling while in 
the air. The genus Arphia, 
also characterized by wings 
with bright red or yellowish 
disks but having the fore wings 
without large spots or blotches, 
usually not even speckled, and 
with the body slenderer than 
in Hippiscus, comprises about 
twenty species scattered over 
the whole country. A. xan- 
thopiera, with plain smoky 
brown fore wings and upper 
body, and hind wings with 
bright yellow disk, broad smoky 
outer band and clearer apex, 
is common in the East; A. 
tenebrosa (Fig. 183), with brown 
and clayey-speckled fore wings 
and upper body and hind 
wings with coral-red disk and 
smoky broad outer band fad- 
ing out in apex, is common 
in the West. The green- 
striped locust, Chortophaga 

Fig. 183. — Arphia tenebrosa. (After Lugger; nat- 
ural size indicated by line.) 

viridijasciata (Figs. 184 and 185), abundant and familiar in the East and 
Mississippi Valley, appears in two forms; in one, the head, thorax, and 

Cockroaches, Locusts, Grasshoppers, and Crickets 145 

Fig. 1S4. — The green-striped locust, Chortophaga viridifasciata, form virginiana, female. 
(After Lugger; natural size indicated by line.) 

Fig. 186. 

Fig. 187. 

Fig. 185. — The green-striped locust, Chortophaga viridifasciata, form virginiana, male. 
(After Lugger; natural size indicated by line.) 

Fig. 186.— The clouded locust, Encoptolophus sordidus, male. (After Lugger; nat- 
ural size indicated by line.) 

Fig. 187. — The pellucid locust, Camnula pelliicida, female. (After Lugger; natural 
size indicated by hne.) 

146 Cockroaches, Locusts, Grasshoppers, and Crickets 

femora are green and there is a broad green stripe on each wing-cover; 
the other form is dusky brown all over; both are about i inch (male) to i^ 
inches (female) long, and have a distinct sharp little median crest on the 

Fig. 190. 

Fig. iqi. 

Fig. 188. — Barren-ground locust, Spharagemon holli, male. (After Lugger; natural size 
of male 20-22 mm., of female 27-33 rnn^-) 

Fig. 189. — Spharagemon collare, race scudderi, male. (After Lugger; natural size in- 
dicated by line.) 

Fig. 190. — The long-horned locust, Psinidia jenestralis, male. (After Lugger; natural 
size indicated by line.) 

Fig. 191. — Circotettix verruculatus, male. (After Lugger; natural size indicated by line.) 

pronotum. The clouded locust, Encoptolophus sordidus (Fig. 186), is another 
species very common in the fall; it is about an inch long, dusky brown 
mottled with darker spots; the wing-covers are blotched and the wings 

Cockroaches, Locusts, Grasshoppers, and Crickets 147 

clear and transparent; the prothorax looked at from above appears to be 
"pinched" at its middle. The males make a loud crackling when in the 

It is famihar knowledge that locusts which are readily seen in the air 
are extremely difficult to distinguish when alighted. This concealment, 
resulting from a harmonizing of the body color with that of the grass or 
soil, is of course an advantage to the locust in its "struggle for existence " 
and is technically known as protective resemblance (see Chapter XVII). No 
locusts show this protective resemblance better 
than the species of Trimerotropis (Fig. 193) 
especially familiar in the western states. The 
colors of various individuals of a single species 
vary with the soil colors of the locality, ranging 
from whitish to 
brownish to slaty 
and bluish. I have 
taken series of spe- 
cimens of Trimero- 
tropis sp. in Colorado 
showing this whole 
range of ground 

Fig. 193. 
(After Lugger; natural size indicated by line.) 

Fig. 192. — Mestohregma cincta, male 

Fig. 193. — The maritime locust, Trimerotropis maritima, female 
ural size indicated by line.) 

(After Lugger; nat- 

The subfamily Tettiginae includes the strange little Acridiids known as 
"grouse-locusts.". They are all under f inch in length, and most of 
them are less than h inch. They have the wing-covers reduced to mere 
scales, but the pronotum is so long that it extends back over the rest of the 

148 Cockroaches, Locusts, Grasshoppers, and Crickets 

thorax to the abdomen and more or less covers it. In some species the 
pronotum actually extends beyond the tip of the abdomen. The head is 
deeply set in the prothorax, the prosternum being expanded into a broad 
border which nearly covers the mouth. As all the grouse-locusts are dark- 
colored and without any conspicuous markings, and choose for habitat the 
dark ground along streams and ponds, or swampy meadows, they are 

Fig. 194. Fig. 105. Fig. 196. 

Fig. 194. — Nomotetlix parvus. (After Lugger; natural size indicated by line.) 
Fig. 195. — Tettigidea lateralis. (After Lugger; natural size indicated by line.) 
Fig. 196. — Tettix granulatus, and pronota of two varieties, (.\fter Lugger; natural size 
indicated by line.) 

infrequently seen except by persistent students. They vary much in colora- 
tion and sHght markings, and harmonize thoroughly with the soil on which 
they habitually live. They feed on lichens, moulds, germinating seeds, 

and sprouting grasses, and are said to eat 
surface mud and muck containing or largely 
consisting of decaying vegetable matter. The 
eggs are laid in a pear-shaped mass in a 
shallow burrow; in May and June the young 
hatch in from sixteen to twenty-five days, 
becoming mature in late fall, or sometimes 
not until the following spring. The nymphs 
and adults hibernate, becoming active again 
early in spring. A common species is Tettix 
gramdatus (Fig. 196), slender, length about 
\ inch, and with the narrow pointed pronotum 
extending beyond the abdomen. This species 
hibernates among rubbish and loose bark, but 
is more or less active on warm winter days. 
It is plentiful all through the rest of the year 
on its feeding-grounds. T. ornatiis (Fig. 197) is a shorter, more robust 
species, and is marked with black spots and indefinite yellow blotches as 

Fig. 197. — Tettix omatus. (After 

Lugger; natural size indicated 

by line.) 
Fig. 198. — Paratettix cucullatus. 

(After Lugger; natural size 

indicated by line.) 

Cockroaches, Locusts, Grasshoppers, and Crickets 149 

indicated in tlie figure. In the genus Tettigidea the antennas have from 
15 to 22 segments, while in Tettix they have only 12 to 14 segments. Tet- 
tigidea lateralis (Fig. 195) is a common species yellovi^ish brown in color, 
more yellowish underneath. It is rather robust and the pronotum extends 
beyond the tip of the abdomen. 

Included in the family Locustidae are katydids, meadow grasshoppers, 
cave-crickets, wingless crickets, western crickets, Jerusalem crickets, and 
what not, but no locusts. The general reader of natural history should 
always keep clearly in mind the 
sharp distinction made by natu- 
ralists between "scientific" and 
"vernacular" names. The ver- 
nacular name locust is applied 
to insects of the family Acri- 
diidae, but not to any of the 
members of the family whose 
scientific name is Locustidae. 
Of the Locustids the best 
known representatives are un- 
doubtedly the katydids. Anna 
Botsford Comstock, the nature- 
study teacher of Cornell Uni- 
versity, introduces them to her 
readers as follows: "The 
chances are that he who lies 
awake of a midsummer night 
must listen, whether he wishes 
to do so or not, to an oft- 
repeated, rasping song that 
says, 'Katy did, Katy did; she 
did, she didn't,' over and over 
again. There is no use of won- 
dering what Katy did or didn't 
do, for no mortal will ever 
know. If, when the dawn 

comes, the listener has eyes r? t, ji ■ j 1 ,■ 1 1 . r • 1 

-' riG. igg. — Broad-winged katydid, and leaf with 

sharp enough to discern one of katydid eggs along edge. (Natural size.) 
these singers among the leaves of some neighboring tree, never a note of 
explanation will he get. The beautiful, finely veined wings folded close 
over the body keep the secret hidden, and the long antenna?, looking like 
threads of living silk, will wave airily above the droll green eyes as much 
as to say, 'Wouldn't you like to know?'" 

150 Cockroaches, Locusts, Grasshoppers, and Crickets 

The katydids are rather large, almost always green insects that live in 
trees and shrubs, where they feed upon the leaves and tender twigs, some- 
times doing considerable injury. With almost all the other Locustids, 
they will also take animal food if accessible, and some of the ground- 
inhabiting forms undoubtedly depend largely on animal substances for 
food. The color and form of the wing-covers and body serve to make them 
nearly indistinguishable in the foliage, and as they do not flock together 
in numbers, they are not frequently seen. Their love-calls or songs, how- 
ever, make the welkin ring at night from 
midsummer until the coming of frost. Few 
katydids sing by day: it would bring their 
enemies, the birds, down on them; but as 
twilight approaches, the males begin their 
shrilling, which is kept up almost constantly 
till daylight. Like the sound-making Acri- 
diids the musical Locustids have a pair of 
special auditory organs, or ears, for hearing 
these love-songs. These ears are tympanal 
organs situated one in the base of each fore 
tibia (the Acridiid ears are on the upper 
part of the first abdominal segment), and 
consist of a thin place in the chitinized 
body-wall (the tympanum), a resonance- 
chamber inside, and a special arrangement 
of nerves and ganglia. There are several 
genera of these Locustids, corresponding to 
the distinctions popularly made under the 
vernacular names narrow-winged, round- 
winged, angular- winged, oblong leaf- winged, 
and broad-winged katydids. The true 
katydid is one of the last-named forms, 
the commonest and most wide-spread species 
being Crytophylliis concavus (Fig. 200). 
It is bright dark-green, and is rarely 
distinguished when at rest in the foliage, although familiar to all from its 
shrill singing. When specimens of katydids are collected and examined, 
concavus may be readily distinguished by the fact that its wings are shorter 
than the wing-covers, and these latter are very convex and so curved around 
the body that their edges meet above and below. The ovipositor of the 
female is short, compressed, slightly curved and pointed. This katydid 
is most in evidence in late summer. People disagree about the melody 
and alleged charm of the song. Many cannot distinguish the "katydid" 

Fig. 200. — Broad-winged katydid, 
CyrtophyUits concavus, male. 
(After Harris; natural size.) 

Cockroaches, Locusts, Grasshoppers, and Crickets 151 

Fig. 201. — Cyrtophyl- 
liis concavus sp. 

syllables, and Scudder, an experienced student of the Orthoptera, says that 

the note, which sounds like xr, has a shocking lack of melody, adding that 

the poets who have sung its praises must have heard 

it at the distance that lends enchantment. The sounds 

are made by the males exclusively, and result from 

the rubbing together of the bases of the wing-covers, 

which have the veins and membrane specially modified 

for this purpose (see Fig. 201). Concavus lays, in the 

autumn, flattened dark slate-colored eggs, about | inch 

long and one-third as wide, in two rows along a twig, 

the eggs overlapping a little. These eggs hatch in the 

following spring, and the young, like the adults, feed 

on the foliage of the tree. 

The oblong leaf-winged and round-winged katydids belong to the genus 

Amblycorypha, and they can be readily recognized by the broad, oblong, 

and rounded wing-covers, and the strongly curved ovipositor of the female, 

with serrated tip. They are grass-green and have the wings longer than 

the wing-covers. The oblong leaf-winged species, A. oblongi folia (Fig. 202), 

is 2 inches long to tips of folded 
wings, while the round - winged 
species, A. rotundijolia, is i\ in- 
ches or less in length. These 
katydids prefer bushes and tall 
weeds or even grass-clumps to 
tree-tops. Oblongijolia is said by 
McNeill to make a "quick shuf- 
fling sound which resembles 
' katy ' or ' katydid ' very slight- 
ly," while the song of rotun- 
dijolia is said by Scudder to be 

made both day and night without variation and to consist of two to four 

notes, sounding like chic-a-chee, run together and repeated generally once 

in about five seconds for an indefinite length of time. 

Fig. 202. — The oblong leaf-winged katydid, 
Amblycorypha oblongijolia, female. (After 
Lugger; natural size.) 

Fig. 203. — Angular-winged katydid, Microcentrum laiirifolium, male. 
(After Riley; natural size.) 

The angular-winged katydids, genus Microcentrum, are large, numerous, 

152 Cockroaches, Locusts, Grasshoppers, and Crickets 

and the most familiarly known of all. The best-known species, M. retinervis, 
is over 2 inches long (from head to tip of folded wings) ; the overlapping 
dorsal parts, of the wing-covers form a conspicuous angle with the lateral 
parts, hence the name "angular-winged." The ovipositor of the female is 
very short, strongly curved, and with a bluntly pointed, finely serrate tip. The 
song of M. laurijoliiim (Fig. 203) is said to sound like tic repeated from 
eight to twenty times, at the rate of four a second. The eggs, of which each 
female lays from 100 to 150 in the fall, are grayish brown, flat, and long- 
oval, about \ inch long by \ inch wide, and are glued in double rows along 
twigs or on the edges of leaves (Fig. 199). I have found them on thorns 
of the honey-locust, and Howard once received "a batch from a western 
correspondent which was found on the edge of a freshly laundried collar 
which had lain for some time in a bureau drawer." The rows are side by 
side, and the fiat eggs overlap each other in their own row. The young 
hatch in spring and, slowly growing, moulting, and developing wings, reach 
full size and maturitv by the middle of the summer. 

Fig. 204a. Fig. 204&. 

Fig. 204a. — The fork-tailed katydid, Scudderia jiircala, female. (After Lugger; nat. size.) 
Fig. 2046. — The fork -tailed haXydid, Scudderia jnrcata,Tadi\e. (After Lugger; nat. size.) 

The narrow-winged katydids, belonging to the genus Scudderia (Figs. 204- 
206), are smaller than the broader-winged kinds, being not more than i^ 
inches in length to tip of folded wing-covers, and the wing-covers are narrow 
and of nearly equal width for their whole length. The ovipositor is broad. 

Fig. 205. Fig. 206. 

Fig. 205. — Scudderia pistillala, female. (After Lugger; natural size ) 
Fig. 206.— Scudderia pistillala, male. (After Lugger; natural size.) 

compressed, and curves sharply upward. These insects frequent shrubbery 
and bushes, or coarse grasses and weeds along ravines or ponds; also 
marshes, cranberry-bogs, and similar wet places. Their flight is noiseless 

Cockroaches, Locusts, Grasshoppers, and Crickets 153 

and zigzag, and when pursued they will take to the lower branches of trees, 
especially oaks if near by. The males sing somewhat in daytime as well 
as at night, and have different calls for the two times. The females lay 
their eggs in the edges of leaves, thrusting them in between the upper and 
lower cuticle by means of their flattened and pointed ovipositor. 

While almost all katydids are green, a few exceptions are known. 
Scudder has found certain pink individuals belonging to a species normally 
green. In mountain regions a few species of gray- or granite-colored katy- 

FiG. 207. — The sword-bearer, Conocephalus ensiger, female. (After Lugger; nat. size.) 

dids are known, the color here being quite as protective as the green of the 
lowland forms, for these mountain species alight to rest on the granite rocks 
of the mountainside. I have found these granite katydids in the Sierra 
Nevada of California. 

Fig. 208. Fig. 209. 

Fig. 208. — The sword-bearer, Conocephalus ensiger, male. (After Lugger; nat. size.) 
Fig. 209. — A common meadow grasshopper, Orchelimum vulgare, female. (After 
Lugger; natural size indicated by line.) 

The meadow grasshoppers are small, katydid-like Locustids, green and 
long-winged, with long, slender hind legs and with the characteristic slender 
thread-like antennae longer than the body. These antennae readily distinguish 
them from any of the locusts (Acridiidae) which may be found in their com- 
pany. The meadow green grasshoppers abound in pastures and meadows. 

154 Cockroaches, Locusts, Grasshoppers, and Crickets 

and they dislike to take to wing, trusting, when alarmed, to spry leaping or 
clever wriggling away and hiding among the lush grasses. Their green 
color of course aids very much in protecting them from enemies. They 

include three common genera, viz.: 
Conocephalus (Figs. 207 and 208), or 
cone-headed grasshoppers or sword- 
bearers with head produced into a 
long, pointed, forward-projecting, cone- 
like process, slender body, and very 
Fig. 210.— a common meadow grasshop- long, slender, straight or angled, sword- 
per, Orchelimum vulgare, male. (After like ovipositor; Orchehmum (Figs. 209 

Lugger: natural size indicated by line.) a \ ^-i ^ ^ j v. 

^^ ^ and 210), the stout meadow grasshop- 

pers, with blunt head, robust body, and short, slightly curved ovipositor; 
and Xiphidium (Fig. 211), the slender or lance-tailed meadow grasshoppers, 
with blunt head, small and slender, graceful body, and nearly straight, 
slender ovipositor, sometimes larger than the body. The eggs of all these 

Fig. 211. — The lance-tailed grasshopper, Xiphidium attenuatiim, female. 
(After Lugger; natural size indicated by line.) 

are laid usually in the stems or root-leaves of grasses, or the pith of twigs. 
The color is usually green, but a few are light reddish brown. The song 
of the males is faint and soft, and is made by day as much as by night. 

Fig. 212. Fig. 213. 

Fig. 212. — Udeopsylla robusta, female. (After Lugger; nat. size indicated by line.) 
Fig. 213. — The spotted wingless grasshopper, Ceuiophilus macidatiis, female. (After 
Lugger; natural size indicated by line.) 

The family Locustidae includes numerous wingless forms, some with 
no remaining trace of wing-covers or wings, some with rudimentary or scale- 

Cockroaches, Locusts, Grasshoppers, and Crickets irr 

like remnants of wing-covers. These latter kinds can sing because the parts 
retained are the sound-producing bases of the wing-covers. The genus 



-Diestrammena mannorata, male; a Japanese locust species found in Minnesota. 
(After Lugger; natural size.) 

Ceuthophilus (Figs. 213 and 215) includes the various species of stone, 
or camel, crickets found all over the country, recognizable by their thick, 

Fig. 2JS.— Ceuthophilus lapidicolus, female. (After Lugger; natural size 
indicated by line.) 

smooth, wholly wingless, brownish body with arched back and head bent 
downwards and backwards between the front legs. They are nocturnal, 

Fig. 216. y^^ _,^^ 

Fig. 216.— The shield-backed grasshopper, Allanlicus pachymerus, male. (After Lue- 

ger; natural size indicated by line.) 
Fig. 217.— The California shield-backed grasshopper, Tropizaspis sp., female. (Nat. size ) 

and during the day hide under stones or logs along streams or in damp woods. 
The individuals of a species which live in the burrows of certain turtles in 
Florida are called "gophers." Perhaps the commonest species, extending 
from New England to the Rocky Mountains, is the "spotted wingless grass- 

156 Cockroaches, Locusts, Grasshoppers, and Crickets 

hopper," C. maculatus (Fig. 213), with sooty brown body dotted with 
pale spots. Some of the wingless Locustids are found in caves, and these 
are either bhnd or have the eyes much reduced. One of these cave-crickets, 
Hadoenuciis siibterraneus, is common in the larger caves of Kentucky, where 
it may be found creeping about on the walls. Garman states that it speedily 
dies when removed from the cave. The genus Atlanticus comprises duU- 

FiG. 218. — The western cricket, Anabrus pttrpurascens, male. (After Lugger; nat. size.) 

colored species with the pronotum extending like a shield back over the 
base of the abdomen, and although the hind wings are wanting, rudimentary 
wing-covers are present, and in the males carry a circular stridulating organ. 

These are called "shield-backed grasshoppers" 
and are to be found in dry upland woods and on 
sloping hillsides with sunny exposure. The two 
common species in the East and the Mississippi 
Valley are .4. dorsalis, with pronotum well rounded 
behind, and A. pachymerus (Fig. 216), with pro- 
notum nearly square. 

A genus similar to Atlanticus found commonly 
in California is Tropizaspis (Fig. 217), the males 


Fig. 219. Fig. 220. 

Fig. 2I(). — The western cricket, Anahriis purpurascens, female. (After Lugger; nat- 
ural size.) 
Fig. 220. — The Jerusalem cricket, Stenopelmahis sp. (Natural size.) 

Cockroaches, Locusts, Grasshoppers, and Crickets 157 

of which sing very pleasantly. In Idaho and other northwestern states 

a large corpulent wingless Locustid, called the western cricket, Anahrus 

piirpurascens (Figs. 218 and 219), often occurs in 

such numbers as to be very destructive to crops. 

The body of this cricket is i^ inches long and 

h inch thick. The ovipositor is three-fourths as 

long as the body, slightly curved, and sword-shaped 

with a sharp point. This species forms march- 
ing armies in Nevada, with two miles of front and 

a thousand feet of depth. On the Pacific Coast 

occurs a large, awkward, thick-legged, transversely 

striped form, Stenopelmatus, called sand-cricket or 

Jerusalem cricket (Fig. 220). It is found under 

stones or in the soil, has a large smooth head with 

"baby-face," and is believed to feed on dead plant 

or animal matter. 

The crickets that we know best are the black 

and brown ones of the house and the fields; but 

there are members of the cricket family, the Gryl- 

lidae, that live in trees and are pale greenish white, fig. 221. -A common 

and others that burrow into the ground and have cricket, Gryllus pennsyl- 

broad shovel-like lore fee,, and still other curious I™; '™»|.l, i^" 

little wingless pygmies that live as guests in ants' dicated by line.) 

nests. But the house- and field-crickets represent the more usual or we 

might say normal and typical kind of Gryllid; 
the others are modifications or ofTshoots of this 
type, both in habit and structure. In all the 
antennae are long and slender (except in the 
burrowing forms, longer than the body), the hind 
legs long and thickened for leaping, and the 
ovipositor, when exserted and visible, long, slender, 
subcylindrical and lance- or spear-like. Well- 
developed wings and wing-covers are present in 
most species, and the males are provided with a 
very effective stridulating organ on the bases of 
the wing-covers. 

In the familiar black, bright-eyed, loud-voiced 
house- and field-crickets the wing-covers when 
folded on the body are flat above and bent down 
sharply at the edge of the body like a box-cover, and the veins in the males 
are curiously changed in course and specially thickened and roughened 
to make a sound-producing organ. This organ is illustrated in Fig. 222. 

Fig. 222. — Cricket and file 
(part of the sound-making 
apparatus). (Cricket nat- 
ural size; the file greatly 

158 Cockroaches, Locusts, Grasshoppers, and Crickets 

To sing, the males lift their wing-covers at an angle of about 45° over the 
back, and strongly rub together the bases. Their chirping is made either 
in the daytime or night, and is a love call or song for their mates. We have 
several common crickets in dwellings, one, Gryllus domesticus (Fig. 223) 
being the European house-cricket, the "cricket on the 
hearth,"' which is becoming at home here, being espe- 
cially met with in Canada. It is pale brown and less 
than an inch long. Gryllus luctiiosus and G. assimilis 
are two native crickets which are common in houses; 
they are black with brownish-black wing-covers, larger 
and nifire robust than domesticus, and with the folded 
wings projecting backward beyond the wing-covers like 
pointed tails. These house-crickets are most active 
at night, and seem to have a taste for almost any 
food-product in the house. They will eat each other 
when other food is scarce. If they become so nu- 
merous in the house that they need to be got rid of, 
advantage may be taken of their liking for sweet 
liquids by exposing smooth-walled vessels half filled 
with such liquids, into which the crickets will fall and 
drown in their attempts to get at the food. The most 
abundant and wide-spread outdoors cricket is Gryllus 
abbreviatus (Fig. 224), the short-winged field-cricket. 
The wings are sometimes wanting, but more often pres- 
ent and shorter than the wing-covers, which in the females are themselves 
unusually short, reaching but half-way to the end of the abdomen. The 
slender ovipositor is as long as the 
body, and the hind femora are ver}- 
thick and have a red spot at the 
base on either side. The life-history 
of this common insect is not yet fully 
known, some writers stating that the 
eggs laid in autumn do not hatch until 
the following spring, the insect thus 
passing the winter in the egg stage, Fig. 224.— The short-winged cricket, Gry/n^ 
while others have taken half-grown abbreviatus. (Natural size.) 

young from beneath logs in late autumn and in midwinter. The field- 
cricket is "nocturnal, omnivorous, and a cannibal. Avoiding the light 
of day," says Blatchley, "he ventures forth as soon as darkness has fallen, 
in search of food, and all appears to be fish which comes to his net. Of 
fruit, vegetables, grass, and carrion he seems equally fond, and does not 

Fig. 223. — The Euro- 
pean house - cricket, 
Gryllus domesticus, 
female. (After Lug- 
ger; natural size in- 
dicated by line.) 

Cockroaches, Locusts, Grasshoppers, and Crickets 159 

hesitate to prey upon a weaker brother when opportunity offers. I have 

often surprised them feasting on the bodies of their com- 
panions; and of about forty imprisoned together in a 

box, at the end of a week but six were Hving. The 

heads, wings, and legs of their dead companions were all 

that remained to show that the weaker had succumbed 

to the stronger — that the fittest, and in this case the 

fattest, had survived in the deadly struggle for existence." 
These crickets hve in cracks in the soil, or under 

stones or logs, or sometimes make burrows. 

The genus Nemobius contains a number of little 

crickets known as "striped ground-crickets," which are 

less than half an inch long, are dusky brownish with hairy 

head and thorax, and have faint blackish longitudinal 

stripes on the head. "Unlike their larger cousins, the 

field-crickets, they do not wait for darkness before seek- 
ing their food, but wherever the grass has been cropped 

short, whether on shaded hillside in the full glare of ^'s^tri ed'^TouTd^! 

the noonday sun along the beaten roadway, mature speci- 
mens may be seen by hundreds during the days of early 

autumn." They are powerful jumpers and readily evade 

attempts to capture them. They feed on living vegetation 

and on all kinds of decaying animal matter, and because of 
their abundance and voracious appetite must do much 
damage at times. Scudder gives the following account of 
the singing of the wingless striped cricket, Nemobius vittatus 
(Fig. 225), our commonest species: "The chirping of the 
striped cricket is very similar to that of the black field-cricket, 
and may be expressed by r-r-r-u, pronounced as though it 
were a French word. The note is trilled forcibly, and lasts 
a variable length of time. One of these insects was once 
The observed while singing to its mate. At first the song was 
mild and frequently broken; afterwards it grew impetuous, 
forcible, and more prolonged; then it decreased in volume 
(Natural size.) ^^^ extent until it became quite soft and feeble. At this 

point the male began to approach 

the female, uttering a series of 
twittering chirps; the female ran 
away, and the male, after a short 
chase, returned to his old haunt, 
singing with the same vigor, but 

cricket, Nemobius 
jasciatus; form vit- 
tatus, female. (After 
Lugger; about 
twice natural size.) 

Fig. 226. — 
snowy tree- 
cricket, CEcan- 
thus niveus. 

Fig. 227. — (Ecanihus jasciatus, female. (After 
Lugger; natural size indicated by line.) 

i6o Cockroaches, Locusts, Grasshoppers, and Crickets 

with more frequent pauses. At length finding all persuasions unavailing, he 
brought his serenade to a close." 

From midsummer till frost comes there is a shrill insistent night-song 
that makes familiar an insect rarely seen except by persistent students. 
X-r-r — r-e-e; t-r-r — r-e-e, repeated without pause or variation about seventy 
times a minute: this is the song of the snowy tree-cricket, or white climbing 
cricket, (Ecantheics niveus (Fig. 226), common all through the East and 
Middle West. These crickets differ much from the better known robust, 
black-brown house- and field-crickets in shape and color; the body is 
about one-half inch long, slender, and the long wing-covers are so held, 
when the insect is at rest, that the back (including the wing-covers) is widest 

behind and tapers forward to the 
small narrow head. The body is 
ivory-white tinged with delicate 
green, and the wing-covers and 
wings are clear. The antennae are 
extremely long and thread-like and 
have two slightly elevated black 
dots on the under side, one on the 
first segment and one on the second. 
The females do much harm by 
their habit of cutting slits in the 
tender canes or shoots of raspberry, 
grape, plum, peach, for their eggs. 
The cane or shoot often breaks off 
at the place where the eggs are 
deposited, and by collecting these 
in the late autumn or winter and 
burning them many eggs will be 
destroyed. Several other species 
of (Ecanthus are found in this 
country; one, O. jasciatiis (Figs. 227 and 228), with three black stripes on 
head and prothorax and usually dark body, is common in the Mississippi 
Valley, and a third species, O. angustipennis, with wing-covers just one- 
third as wide in broadest part as their length, is less common. 

Occasionally one finds on the ground, or more likely in digging, a curious 
flattened, light velvety brown insect about an inch and a half long, with 
the fore feet much widened and strangely resembling those of the common 
mole, and altogether having an appearance strange and unlike that of any 
other insect. This is a burrowing, or mole, cricket, which burrows beneath 
the soil in search of such food as the tender roots of plants, earthworms, 
and the larvae of various insects. Its eyes are also like those of the mole, 

Fig. 228. Fig. 229. 

Fig. 228. — (Ecanthus fasciati4S, rrnde. (After 

Lugger; natural size indicated by line.) 
Fig. 229. — Orocharis saltator, fermle. (After 

Lugger; natural size indicated by line.) 

Cockroaches, Locusts, Grasshoppers, and Crickets 1 6 1 

much reduced, being nearly lost, and as this cricket crawls rather than 
leaps, the hind or leaping legs are not so disproportionately larger than the 
others as in the above-ground crickets. The males make a sharp chirping 
loud enough to be heard several rods away. The common species, called 
the northern mole-cricket, Gryllotalpa borealis, has the wing-covers less than 
half the length of the abdomen, while the wings extend 
only about one-sixth of an inch beyond them. A less 
common species, G. Columbia, the long- winged mole- 
cricket, has the hind wings extending beyond the 
tip of the abdomen. The mole-crickets like rather 
damp places near ponds or streams, where they make 
channels with raised ridges which resemble miniature 
mole-hills. These "runs" usually end beneath a stone 
or small stick. The insects are infrequently seen, as 
they remain mostly underground, only occasionally 
coming out at night. The female deposits from two 
hundred to three hundred eggs in masses of from 
forty to sixty in underground chambers, and the young 
are about three years in reaching maturity. When 
present in any region in large numbers mole-crickets Fig. 230. — The Porto 

become seriously destructive because of their attacks ^ican molc-cncket, 

bcapteriscus didacty- 
on plant-roots. In Porto Rico a mole-cricket, Scap- lus. (After Barrett; 

teriscus didadylus (Fig. 230), called "changa," dam- natural size.) 

ages tobacco, sugar-cane, and small crops to the value of more than 

$100,000 annually and is by far the most serious insect 

pest in the island. 

Much smaller than the true mole-crickets are the 

pygmy, burrowing crickets of the genus Tridactylus, 

of which several species occur in the United States. 

The largest species, T. apicalis (Fig. 231), is about 

J inch long. They resemble the mole-crickets in general 

body characters, but are more brightly colored, and the 

fore feet, although broad and flat for digging, differ in 

being curiously armed at the end with three spurs; hence 

Fig. 27,1 —Tridactylus ^]^g oreneric name. They can leap amazingly, so that 
apicahs. (After'' . . ■' ^ ^^' 

Lugger; natural size they seem, on jumping, to disappear most mysteriously, 

indicated by line.) the eye not being able to follow them in the air. 
The most aberrant of all the crickets are the tiny flat and broad-bodied 
species of the genus Myrmecophila, which live as commensals or mess- 
mates in the nests of ants. They are found only in ants' nests, have no 
compound eyes, and the hind femora are much swollen and enlarged. 

i62 Cockroaches, Locusts, Grasshoppers, and Crickets 

The semi-parasitic life which they lead has resulted in such a change of 
habits that their body is modified very far from the normal cricket type. 
The commonest species is Myrmecophila nebrascensis, about -j\ inch long, 
shown in Fig. 232. 

Formerly included in the order Orthoptera, the earwigs are now recog- 
nized as entitled to distinct ordinal rank, and the thirty or more genera in 
the world, of which but six occur in the United States, 
are held to constitute the order Euplexoptera. This 
order is closely related to the Orthoptera, although the 
insects themselves look more like beetles. 

The earwigs are small, brownish or blackish insects, 
readily recognized by the curious forceps-like appendages 
on the tip of the abdomen (Fig. 233). They are either 
winged or wingless, but when winged have small leath- 
ery wing-covers only extending about half-way to the 
tip of the abdomen, with the well-developed nearly 

T?Tp 2 2 2 A^ V f fft C" 

cophila nebrascensis, a hemispherical wings compactly folded, both longitudi- 
degenerate cricket nally and transversely, underneath them. Earwigs are 
times "^ot often seen because they are nocturnal in habit, 
but in some places they are rather abundant. They 
are ve<^etable feeders, being especially fond of ripe fruit, flower corollas, etc., 
which they bite off and chew with the well-developed jaws and maxilla?. 
The female lays her small, yellowish oval eggs in 
small masses under fallen leaves or in other con- 
cealed places, and is said to nestle on them as a 
hen on her eggs. She is also said to protect the 
young for some time after they are hatched. The 
young undergo an incomplete metamorphosis, de- 
veloping wings externally, and resembling the 
parents, except in size, from the time of their 

The commonest representative of the order in the 
northern and eastern states is the little earwig. 
Labia minor (Fig. 233), measuring to tip of forceps only about J inch. 
Other American species, as Labidura riparia, a Florida species, brownish 
yellow with a pair of longitudinal black stripes on prothorax and wing- 
covers, with long slender forceps, and Anisolabis annulipes, a black wingless 
California species with short heavy forceps, are larger, these two species 
being a Httle more and a little less than | inch respectively. 

that inhabits 
nests. (Five 
natural size.) 

Fig. 233. — An earwig, 
Labia minor. (Six times 
natural size.) 


INSECTS, ETC. (Order Hemiptera), AND THE 
THRIPS (Order Thysanoptera) 


HEN an Englishman says "bug" — and 
he doesn't say it in polite society — he 
means that particular sort of bug which 
we more specifically speak of as bedbug; 
when we say "bug" we are likely to mean any insect 
of any order; but when a professed student of insects, 
an entomologist, says or writes bug, he means some 
member of the insect order Hemiptera. It is to this 
order of "bugs" that we have now come in our system- 
atic consideration of insects, and it is in this order that 
we first meet conspicuously the difficulties of treating 
systematically the populous insect class. From now 
on the making of this book useful depends on the discriminating selection 
of the few kinds of insects whose special consideration the limits of 
text and illustration permit, leaving the great majority of species to be 
referred to comprehensively and vaguely as the "others." 

The Hemiptera, or true bugs, make up a large order compared with any 
of those so far considered, although a smaller one than certain others yet to be 
taken up. As regards popular acquaintanceship and interest also this 
order is still more inferior to the other large ones, namely, the beetles, the 
moths and butterflies, the two-winged flies, and the ants, bees, and wasps. 
Most of the true bugs are small, and obscurely, or at least inconspicuously, 
colored, and few of them attract that attention necessary to gain popular 

The order Hemiptera includes over 5000 known species of North 
American insects, representing a large variety and a great economic impor- 
tance; some of the most destructive crop pests and most discomforting insect- 
scourges of man and the domestic animals belong to this order. The 
chinch-bug's lavages in the corn- and wheat-fields of the Mississippi Valley 
offer effective evidence to the dismayed farmers of the workings of a dis- 
pleased Providence; the tiny sap-sucking aphids and phylloxera and insig- 




164 Bugs, Cicadas, Aphids, and Scale-insects 

nificant-looking scale-insects make the orchardist and vine-grower similar 
believers in supernatural moral correction by means of insect -scourges, 
and the piercing and sucking lice and bugs — in the English meaning — make 
personal and domestic cleanliness a virtue that brings its own immediate 

Other not unfamiliar representatives of this order are the loud-singing 
cicadas with their extraordinarily protracted adolescence, the thin-legged 
water-striders and skaters of the surface of pond and quiet trout-pool, the 
oar-legged water-boatmen and back-swimmers of the depths of the same 
pools, the ill-smelling squash-bugs, calico-backs, and stink-bugs of the 
kitchen-gardens, the big, flat-bodied, electric-light or giant water-bugs that 
, whirl like bats around the outdoor arc-lights, 

and the assassin- and "kissing "-bugs of one- 
time newspaper interest. In structure all the 
Hemiptera agree in having the mouth-parts 
formed into a piercing and sucking beak (Fig. 
234) capable of taking only liquid food. As 
that food is nearly always the blood of living 
animals or the sap of living plants, the nearly 
uniformly injurious or distressing character of 
the food-habits of all the members of the 
order is apparent. This beak is composed 
of the elongate, tubular under-lip (labium) 
acting as sheath for the four slender, needle- 
like piercing stylets (modified mandibles and 
maxillae). The labium is not a perfect tube, 
for it is narrowly open all along its dorso- 
medial line, but the edges of this slit can be 
brought closely together and the slit also 
covered internally by the stylets, so that an 
effective tubular sucking proboscis is formed 
(Fig. 14). The name Hemiptera is derived from 
the character of the fore wings shown by most, 
though by no means all, of the members of the 
order; this is the thickening of the basal half of the otherwise thin, 
membranous wing, so that each fore wing is made up of two about equal 
parts of obviously different texture and appearance; hence "half-winged" 
(Fig. 268). All Hemiptera (excepting the male scale-insects) have an incom- 
plete metamorphosis, the young at birth resembling the parents in most essen- 
tial characteristics except size and the presence of wings. By steady growth, 
with repeated moultings and the gradual development of external wing- 
pads, the adult form is reached, without any of the marked changes apparent 

Fig. 234. — Diagram of section 
through head and basal part 
of beak of a sucking-bug. 
ph., pharynx; m., muscles 
from pharynx to dorsal wall 
of head; •y., valve; 5., stop- 
per; m., muscle of stopper; 
s.d., salivary duct; Ir., la- 
brum; b., one of the stylets 
of beak. To pump fluid up 
through the beak, the mus- 
cle attached to the stopper 
contracts, thus expanding the 
cavity closed by the valve. 
(After Leon.) 

Bugs, Cicadas, Aphids, and Scale-insects 165 

in the insects of complete metamorphosis. With similar mouth-parts the 
young have, in most cases, similar feeding habits, preying on the same kinds 
of plants or animals that give nourishment to the parents. 

The extent of the injuries done by various members of this order to 
farm and orchard crops, to meadov^s and forests, and to our domestic 
animals is enormous. Of the other insects the order of beetles includes 
numerous crop pests, and the caterpillars of many moths and a few butter- 
flies do much damage; locusts have a healthy appetite for green things, 
and many kinds of flies could be lost to the v^^orld to our advantage, but 
perhaps no other order of insects has so large a proportion of its members 
in the category of insect pests. The single Hemipterous species, Blissus 
hiicopterus, better known by its vernacular name of chinch-bug, causes 
an annual loss to grain of twenty millions of dollars; the grape phylloxera 
destroyed the vines on 3,000,000 acres of France's choice vineyards; the 
San Jose scale has in the last ten years spread from California to every 
other state and territory of the United States and become a menace to the 
whole fruit-growing industry. So, despite their small size and their general 
unfamiliarity to laymen, the Hemiptera are found by economic entomologists, 
in their warfare against the insect-scourges of the country, to be one of the 
most formidable of all the insect orders. 

The classification of the Hemiptera into subgroups is a matter Hkely 
to prove difficult for the amateur and general collector. The order as repre- 
sented in our country includes thirty-nine families, and the structural char- 
acters separating some of these families are slight and not easily made out by 
untrained students. For the use, however, of readers of this book capable 
of using them, keys or tables of all the families of the Hemiptera are presented. 
For more general use, however, I shall try to arrange the families in groups 
depending on the habits and more obvious appearance and make-up of the 
insects, characteristics which may be readily noted. And this arrange- 
ment will not be less "scientific" than the arrangement in the key com- 
monly used by entomologists, as the latter is confessedly largely artificial 
and convenient rather than natural in its groupings. 

The order is separable into three primary natural groups or sub-orders 
as follows: 

Wingless forms, with a fleshy, unsegmentcd sucking-beak, living as parasites on 

man and other mammals Parasita. 

Winged, or sometimes wingless, but always with the beak segmented. 

Wings of the same texture throughout and usually held sloping or roof-like 
over the back and sides of the body; sucking-beak arising from the 
hinder part of the lower side of the head; tne head so closely joined 
to the prothorax that the bases of the fore legs touch the sides of the 
head Homoptera. 

1 66 Bugs, Cicadas, Aphids, and Scale-insects 

Fore wings with basal half thickened and parchment-like, apical half thin 
and membranous; the four wings lying flat on the back when folded, 
the membranous tips overlapping; sucking-beak arising from the 
front part of head, and the head usually separated from the pro- 
thorax by a m.ore or less distinct neck Heteroptera. 

Of these three suborders the Parasita, or sucking-Hce, are degenerate 
wingless species and will be considered last. The Heteroptera include 
the so-called "true bugs" with fore wings thickened at base, and when 
folded lying fiat on the back, as the squash-bug, chinch-bugs, and the great 
majority of the species in the order, while the Homoptera include the cicadas, 
the tree- and leaf-hoppers, the aphids or plant-lice, the mealy-winged flies, 
and the degenerate scale-insects. 


Key to Families of the Homoptera (includes both Nymphs and Adults). 

(Adapted from Woodworth.) * 

Proboscis seeming to rise from the middle of the sternum, or proboscis wanting; insects 
less than J inch long. 

Hind femora much larger than other femora (Jumping plant-lice) Psyllid.e. 

Hind femora not much larger than the others. 

Legs long and slender (Plant-lice.) Aphidiid.?;. 

Legs short, or wanting. 

Feet of one joint, or wanting (Scale-insects.) Coccid^. 

Feet of two joints (Mealy wings.) Aleyrodid.^:. 

Proboscis plainly arising from the head. 

With three ocelli, sometimes (nymphs) with large front tibiae and no wings. 

(Cicadas.) Cicadid^. 
With two ocelli or none, and the front tibiae not enlarged. 

Antennae inserted on head below the eyes (Lantern-flies.) Fulgorid.e. 

Antennae inserted in front of and between the eyes. 

Prothorax extending back over the abdomen (Tree-hoppers) Membracid^. ^ 

Prothorax not extending back over the abdomen. 

Hind tibiae with few spines (Spittle-insects.) CERCOPiD.aE. <3 

Hind tibse with two rows of spines (Leaf-hoppers.) Jassid^. ^ 

Perhaps no other insect-species has any single characteristic of its hfe- 
history of the same interest as the extraordinarily long duration of the adoles- 
cence of the seventeen-year cicada. That a single one of the 300,000 and 
more known species of insects should have a period of development from 
egg to adult of more than sixteen years, while this period in all other insects 
varies from a few days to not more than three years — comparatively few 
insects live, all told, more than a year — is perhaps the most striking excep- 
tional fact in all insect biology. The other members of the family 
Cicadidae, to which this insect belongs, have, as far as known, an immature 

Bugs, Cicadas, Aphids, and Scale-insects 167 

life of but one or two years. But few species of cicadas, dog-day locusts, 
harvest-flies, or lyremen, as they are variously called, occur in this country 
— they are more abundant in subtropic and tropic countries — but their 
large, robust, blunt-headed body, their shrill singing and their wide dis- 
tribution make them familiar insects. 
In summer and fall the piercing, 
rhythmic buzzing of the cicadas comes 
from the trees from early morning 
till twilight. The song, unlike that 
of the katydids and tree-crickets, is 
hushed at night. The sound is made, 
not by a rasping together of wings 
or legs, but by stretching and relaxing 
a pair of corrugated tympana, or 
parchment-like membranes, by means 
of a muscle attached to the center 
of each; much, indeed, as a small 

boy makes music from the bottom of Fig. 235. — The seventeen-year cicada, 
.,, . • r . J J. -. Cicada septendecim; specimen at left 

a tm pan with a strmg fastened to its growing sound-making organ, v.p., 
center. These sound-making organs ventral plate; /., tympanum. (Natural 
of the cicadas, confined to the males — ^^^^'' 

"Happy is the cicada, since its wife has no voice," says Xenarchos — are 
situated in resonance-cavities or open boxes, furnished with other sym- 
pathetically vibrating membranes, at the base of the abdomen (Figs. 235 
and 236). The sound-chambers are incompletely closed (wholly open in the 

seventeen-year cicada) by a pair of semicircular 
disks, which are opened or shut by move- 
ments of the body so as to give the song a 
peculiar rhythmic increase and decrease of 

The cicada that is most familiar, and.on hand 
every summer over most of the country, is the 
large (2 inches in length to tip of closed wings) 
black and green dog-day harvest-fly, Cicada 
tibicen. The life-history of this species is not 
fully known, but the insect requires, accord- 
ing to Comstock, two years to t^ecome mature. 
The really famous cicada is Cicada septen- 
decim, the seventeen-year locust, or periodical cicada (Fig. 235). It is 
about li inches long, black, banded with red on the abdomen, and with 
bright red eyes and the veins of both wings red at the base and along the 
front margin. The females lay their eggs in early summer in slits which 

Tig. 236. — Diagram of section 
of body of cicada, showing 
attachment of muscles to inner 
surface of sound-making 
organ. (Enlarged.) 

I 68 Bugs, Cicadas, Aphids, and Scale-insects 

they cut with the sharp ovipositor in the twigs of various trees, in this way 
often doing much damage to orchards and nurseries. The young hatch 
in about six weeks and drop to the ground, where they burrow down through 
cracks and begin their long underground life. They feed on the humus 
in the soil and, to some extent, on juices sucked from the tree-roots. They 
grow slowly, moulting probably four or six times at intervals of from 
two years to four years. In spring or early summer of the seventeenth 
year (thirteenth in a race in the southern states) they come above ground, 
and, after hiding for a while under stones and sticks, crawl up on the trunks 
of trees and there moult for the last time, the winged adult emerging and 
soon flying into the tree-tops. The various broods or swarms in this country, 
about twenty in number, are known, and the territory occupied by each 
has been mapped, so that it is possible for entomologists to predict the 
appearance of a swarm of seventeen-year cicadas in a particular locality 
at a particular time. As all the members of one of these swarms issue in 
the same season, and indeed in the same month or fortnight, they usually 
attract much attention. The broods to issue in the next few years are the 
following: a large one in 1905 in the northern half of Illinois, eastern part 
of Iowa, southern part of Wisconsin, southern edge of Michigan, and northern 
and western edge of Indiana; a scattered one in igo6 ranging, not contin- 
uously, from Massachusetts south and west through Long Island, New 
Jersey, Pennsylvania, Maryland, West Virginia, Ohio, Indiana, Kentucky, 
Tennessee, North and South Carolina, and northern Georgia; and a large 
one in 1907, ranging from central Illinois south and east to the Gulf and 

A considerable number of small insects, often seed-like in shape, or 
with the thorax prolonged into odd horns, spines, or crests, are included 
in the families of tree-hoppers (Membracidae) and lantern-flies (Fulgoridse) 

(Fig. 237). Striking members, large and bright- 
colored, of this latter family are found in the 
South American tropics, but the North American 
species are small, and are rarely seen or collected 

FiG_ 237. A fulgorid, Stohera by amateurs. Among the commonest of our forms 

tricarinata. (After Forbes; are the candle-heads, species of Scolops, small 

natural length 1 inch.) . . t • j u u -i-i. 4.u u j 

msects livmg on grass and herbage, with the head 
bearing a long slender upcurving projection. The tree-hoppers (Mem- 
bracidae) almost all suggest small angular brownish seeds or thorns in shape 
and color. The prothorax is sometimes widely expanded, sometimes 
lengthened so as to cover nearly the whole body, sometimes humped or 
crested, sometimes spined or pitted. The unusual form is probably pro- 
tective, making the insects simulate seeds or other plant structures. The 
species of Enchenopa (Fig. 239) are curiously horned. E. hinotata is common 

Bugs, Cicadas, Aphids, and Scale-insects 169 

in the east. It is gregarious and is attended by ants which feed on a sweetish 
substance excreted by it. It lays its eggs in Httle white waxen frothy masses. 
A curiously humpbacked form is Senilia camelas (Fig. 240). The best known 

and most injurious tree-hoppers are those 
of the genus Cerasa, of which the species 
^C. bubalus, or buffalo tree-hopper (see initial 
letter of this chapter), injures fruit-trees 
both by piercing and sucking sap from 


::■ ,nw^ 

Fig. 238. Fig. 239. Fig. 240. 

^ Fig. 238. — The black-backed tree-hopper, Arthasia galleata. (After Lugger; natural 

length I inch.) 
' Fig. 239. — A tree-hopper, Enchenopa gracilis. (Three times natural size.) 
^ Fig. 240. — A tree-hopper, Senilia camelas. (Three times natural size.) 

them, and by making slits in the twigs to lay eggs in. It is about ^ inch 
long, light grass-green with whitish dots and a pale yellowish streak on 
each side. On the front there are two small sharp processes jutting out one 
on each side from the prothorax, and suggesting a pair of horns, hence 
the name. It is common on apple and many other trees from the middle 
of summer until late in the autumn. The eggs are laid in pairs of nearly 
parallel and slightly curved slits. The young hatch in the spring following 

Walking over our lawns or through pastures and meadows we often 
startle from the grass hundreds of small, usually greenish, little insects that 
leap or fly for a short distance, but soon settle again in the herbage. Nearly 
all these smiiU and active insects are sap-sucking leaf-hoppers, of the family 
Jassidas, one of the largest and most injurious of the Hemipterous families. 
It is stated by careful students of these grass-pests that from nearly one- 
fourth to one-half of all the grass springing up annually is destroyed by 
leaf-hoppers. Professor Osborn estimates that over one million leaf-hoppers 
can and often do live on an acre of grass-covered ground. These insects 
are rarely more than J inch long, and most of them are nearer half of that. 
The body is more slender than in the tree-hoppers, and is usually widest 
across the prothorax or a Httle behind it, tapering back to the tip of the 
folded wings. The head is more or less triangular, as seen from above, 
and the face is oblique, sloping back to the base of the fore legs. The 
family is a large one, containing many species, of which several are well 

170 Bugs, Cicadas, Aphids, and Scale-insects 

known to economic entomologists as special pests of grasses, growing grain, 
grapes, roses, etc. The injury is caused by the draining away of the 
sap of the plant by the host of little sucking-beaks thrust into its leaves 
or stem. Among the notorious destructive species are the destructive leaf- 
hopper,"^ CicaJn/a exitiosa, i inch long, brownish, which often injures 
seriously the winter wheat of the southern states. Also the various 
grape-leaf hoppers, which cause the leaves of grape-vines to wilt and turn 
brown and prevent the formation of full grapes; one 
of these, Erythroneura vitis, is about ^ inch long, 
crossed by two blood-red bands and a third dusky 
one at the apex. I have seen millions of individuals 
oi^Erythroneura comes (Fig. 242) in the great 3300- 
acre vineyard of the Vina Ranch in the Sacramento 
Valley of California. These leaf-hoppers hibernate 
in the vineyard or about its edges under fallen 
leaves and rubbish. Probably the best remedy for 
them is to keep the vineyards as clean as possible, 
.1 or at least to burn up in the winter any accumulated 

rubbish. The rose leaf-hop- 
per, ' Tr^/z/or^'Zja rosa, is 
often abundant on rose- 
bushes, and also on apple- 
trees. The eggs are laid in 
the summer, and the young 
develop through the summer 
and fall, hibernating as 
adults under leaves or rub- 

A common leaf-hop- 

FiG. 241. Fig. 242. 

Fig. 241. — The celery leaf -hopper, Cicadula 4-Uneata. 

(After Lugger; natural size indicated by line.) 
Fig. 242. — Two vine-hoppers, at left Erythroneura _ 

viilnerata, on right^£. comes. (After Forbes; much per of grass-fields is' Diedro- 

en arge . j cephala molHpes, ^ inch long, 

spindle-shaped, grass-green above, pale yellowish below, with black lines 

across the face and top of head, and the fore wings with bluish veins and 

yellowish edges. 

Occasionally one finds frothy, spittle-like masses adhering to the stems 
of weeds or shrubs in which may be found imbedded one or more odd- 
shaped, squat, slant-faced insects from -j\ inch to ^ inch long (Fig. 243). 
These are the young — they have no wings, only wing-pads or, if very young, 
not even these — of the spittle-insects or frog-hoppers, family Cercopidae. 
The spittle is a viscid fluid expelled from the alimentary canal of the insects, 
and beaten up into a froth by the whisking about of the body. What 
advantage it is to the young insects is hard even to conjecture; it certainly is 
not known. The adult frog-hoppers — this name is derived from a popular 

Bugs, Cicadas, Aphids, and Scale-insects 171 

belief that the spittle is that of tree-frogs — are small flattish, brownish or 

grayish insects about -3- inch long which occasionally occur in sufficient 

numbers to do some injury to grapes, 

cranberries, or pasture grasses. A grape 

frog - hopper, ^yl/j/^ro/'/zora ^-notata, has 

brown wing-covers with three blackish 

spots on each; another found on grapes 

in the east, A. signoreti, is tawny brown 

clouded with dull white and thickly dotted 

with black spots; the cranberry spittle- 

insecf; Clastoptera protens, which occurs 

on cranberries and blueberries in marshes, 

is black, with two yellow bands on top 

of the head, one in the thorax, two 

oblique stripes on the base of the fore 

wings, and a cross-bar near the tip; C. 

pini is a small shining black species with 

pale yellow head with black band at front 

margin, that occurs on the needles of 


Looking like miniature cicadas, but 
belonging to a diiTerent family, and really 
more nearly related to the aphids or true 
plant-lice, are the Psyllidae, or jumping 
plant-lice. They are not more than \ inch long, their hind legs are 
enlarged for leaping, some of them exude honey-dew, as the true plant- 
lice and the scale-insects do, and some make galls on the wings of hack- 
berry and other trees. The best-known and most destructive member of 
the family is the pear-tree flea-louse, Psylla pyricola. This is a minute 
insect measuring only -^-^ inch long to tip of folded wings, but it often occurs 
in such large numbers in pear-orchards in the northeastern and northern 
states as to destroy extensive orchards. The eggs are orange-yellow and 
laid on the leaves, each egg having a lash-like process projecting from it. 
The young is broad and flat and yellow in color, growing brownish as it 
grows older. The adults hibernate in crevices in the bark and come out 
in spring to lay their eggs. The pests can be killed by spraying the trees 
with kerosene emulsion (see p. 189), immediately after the leaves have 
expanded in the spring. 

A very important and very interesting family is that of the Aphidiidae, 
the plant-lice or aphis-flies (Figs. 244 and 245). The species, of which 
there are many, are all small, \ inch being a rarely attained maximum 
length. The most familiar representatives of the family are the tiny, 


Fig. 243. — The spittle-insect, ApJiro- 
phora, showing stages of froth 
production. (After Morse; en- 

172 Bugs, Cicadas, Aphids, and Scale-insects 

^ __ are 

Fig. 244;-The southern grain plant- ^f ^^ plant-lice which infest 

louse, loxoptera gramineum, winged ^ 
migrant. (After 

plump-bodied, pale-green insects, some with two pairs of long, delicate, 
transparent wings, some without wings, common on flowers in conserva- 
tories and gardens and known as "green fly." Other often-noticed kinds 
are the cockscomb gall-louse of the elm and the "blights" of various foliage 

trees, as alder-blight, beech-blight, elm- 
blight, etc., these "blight" aphids all 
secreting conspicuous white woolly masses 
of wax and most of them also excreting 
honey-dew, which is conspicuous on the 
leaves and on the sidewalks under the 

Of more economic importance 

Pergande; much crop-plants, the extraordinarily ruinous 
enlarged.) grape-phylloxera, for example, the apple- 

tree, and the woolly apple-aphis, the cherry-, plum-, and 
peach-aphids, the corn-root louse, the hop-louse, and the cabbage-aphis, 
turnip-louse, and other aphid pests of garden vegetables. All of these 
insects are minute soft-bodied defenceless creatures, which effect their great 
injuries to their host-plants by virtue 
of great numbers. Fitch, New York's 
first state entomologist, estimated the 
number of cherry-aphids that were 
living at one time on a small young 
cherry-tree to be 12,000,000. Although 
uncounted millions of the toothsome 
juicy little aphid bodies are being con- 
stantly eaten in spring and summer by 
eager predaceous insects, such as lady- 
bird beetles, certain syrphid-fly larva; 
and aphis-lions (larvae of lace-wing and hemerobius flies), just as constantly 
are new millions being produced by the fecund aphis mothers, most of the 
young being born alive and requiring but a few days to complete their 
growth and development, and to be ready to take up the production of 
young themselves. 

Professor Forbes has made an estimate of the rate of increase of the corn- 
root louse that shows this great fecundity. A single stem-mother of the 
corn-root aphis produces twelve to fifteen young that mature in a fortnight. 
"Supposing that all the plant-lice descending from a single female hatched 
from the egg in spring were to live and reproduce throughout the year, we 
should have coming from the egg the following spring nine and a half tril- 
lion young. As each plant-louse measures about 1.4 mm. in length and .93 

Fig. 245. — The southern grain-louse, 
Toxoplera gramineum, wingless. A, 
female; B, young nymph; C, older 
nymph. (After Pergande; much en- 

Bugs, Cicadas, Aphids, and Scale-insects 173 

mm. in width, an easy calculation shows that these conceivably possil)le de 
scendants of a single female would, if closely placed end to end, form a proces' 
sion seven million eight hundred and fifty thousand 
miles in length; or they would make a belt or strip 
ten feet wide and two hundred and thirty miles long." 
The remarkable plasticity of the aphids as re- 
gards their possession or lack of wings and, on the 
physiological side, their reproduction agamically or 
sexually, introduces certain unusual conditions into 
their life-history. Although each species is likely 
to present idiosyncrasies of its own, a fair example 
of the course of aphid life through a season may 
be outlined as follows: In spring there hatch, from 
eggs which have been laid the fall before, wingless 
females, called stem-mothers, which produce young 
agamically (i.e., from unfertilized eggs) either by 
giving birth to them in active free condition or by 
laying eggs. From these eggs hatch wingless females 
which produce in turn other agamic broods of wing- 
less females. But at any time in the course of these 
successive agamic generations either all or a part of 
the individuals of a brood may be winged, and these 
winged females fly away to other plants and there 
found new colonies which continue the series of 
agamic generations. But toward the end of the 
season, when the first cold weather announces the 
approaching winter, broods, still parthenogenetically 
produced, of sexed individuals, both males and fe- 
males appear. "The males may be either winged 
or wingless, but these true females are always wing- 
less." These individuals mate, and each female 
produces a single large egg which passes over 
the winter to give birth in the following spring to a 
wingless stem-mother — that one which begins the 
spring series of parthenogenetic generations. The unfertilized eggs, called 
pseud-ova, produced in numbers by the spring and summer agamic mothers 
(from which eggs the young frequently emerge while the eggs are still in the 
body of the mother) should not be confused with the single fertilized egg 
laid in the late fall by the mated females of the sexed generation. Although 
these two sorts of eggs are alike in their earliest stages in the ovaries of the 
females, differences very soon occur, the embryo in the pseud-ovum begin- 
ning to develop before the formation of its own egg is properly completed. 

Fig. 246. — Bodies of 
aphids which have been 
killed by Hymenoptcr- 
ous parasites, the adult 
parasitic flies having 
emerged from the small 
circular holes. (En- 

174 Bugs, Cicadas, Aphids, and Scale-insects 

Characteristic variations in the general course are described later in 
connection with the accounts of the few particular aphid species for which 
we have place, but it should be kept in mind that considerable variations 
may occur in the case of a single species. Extrinsic influences, such as 
crowding a host-plant and hence the lessening of food, or an unusual 
humidity or lack of humidity, an early lowering of temperature in autumn, 
etc., seem to be very potent in producing or acting as effective stimuli for 
adaptive variations of the usual course of life. Slingerland reared ninety- 
four successive generations (in 
four years) of an aphid species 
in the insectary at Cornell 
University under such constant 
conditions of food-supply and 
summer temperature that not 
a single winged aphid nor single 
sexual generation was pro- 
duced. Even longer series of 
identical wingless agamic gen- 
erations have been obtained by 
certain European experiment- 
ers. Clarke, in California, has 
been able to produce a winged 
generation at will by simply 
changing the chemical constitu- 
tion of the sap of the host- 
plant on which the aphids were 
reared in his laboratory. 

In addition to the interest- 
ing variation as regards wings 
and reproductive processes 
among the various individuals 
of a single aphid species, it has been found that of the wingless males some 
have no mouth, while others are furnished with functional mouth-parts 
and opening. An interesting physiological variation also occurs in the 
matter of the food-plant selected. The winged individuals frequently 
migrate to a plant of different species from that on which they were born. 
For instance, the apple-aphids. Aphis mali, "spend the summer upon grasses, 
where they continue breeding until autumn, when they return to the apple and 
the winged females establish colonies of the wingless egg-laying form upon 
the leaves. The males fly in from the summer host-plant; the eggs are 
then laid on the twigs and buds, and the cycle for the year is completed." 
The common cherry-aphis, Myzus cerasi, has a similar history, described by 

Fig. 247. — Rose-aphids visited by ants, 
size; from life.) 


Bugs, Cicadas, Aphids, and Scale-insects 175 

Weed as follows: "It winters over on the twigs in the egg state. Early 
in spring the young aphids hatch and crawl upon the bursting buds, insert- 
ing their tiny sap-sucking beaks into the tissues ~of the unfolding leaves. 
In a week or ten days they become full-grown and begin giving birth to 
young lice, that also soon develop and repeat the process, increasing very 
rapidly. Most of the early spring forms are wingless, but during June 
great numbers of the winged lice appear, and late in June or early in July 
they generally leave the cherry, migrating to some other plant, although 
we do not yet know what that plant is. Here they continue developing 
throughout the summer, and in autumn a winged brood again appears and 
migrates back to cherry. These migrants give birth to young that develop 
into egg-laying females which deposit small, oval, shining black eggs upon 
the twigs." 

The point of all this is plainly that in the aphids there must be recog- 
nized an unusual and, to them, very advantageous adaptive plasticity of both 
structure and function. Defenceless as are the aphid individuals as far 
as capacity either to fight or to run away is concerned, the various aphid 
species are, on the contrary, very well defended by their structural and physi- 
ological plasticity and their extraordinary fecundity. 

The two secretions, wax and honey-dew, play an important part in the 
aphid life. The wax secreted or excreted through various small openings 
scattered over the body is, of course, liquid when first produced, but quickly 
hardens; the total waxy secretion appears usually as a mass of felted threads 
or "wool," and doubtless is an important protection for the deUcate body. 
The honey-dew, long supposed to be secreted through two conspicuous 
tubular processes on the dorsal surface of the posterior end of the abdomen, 
is now known to be an excretion from the intestine, issuing in fine droplets 
or even spray from the anal opening. From the so-called "honey-tubes" 
issues another secretion, not sweetish, about which Httle is known. It is 
common knowledge, however, that the aphid honey-dew is a favorite food 
of ants— the Germans call it the ants' "national dish"— and many accounts 
have been written of the care of plant-lice, the ants' cattle, by the ants them- 
selves. Without question there is some basis of fact for these stories. No 
more evidence of this is needed than the careful observations of Professor 
Forbes of the extraordinary care of the corn-root louse by the little brown 
ant, Lasius brnnneus, of the Mississippi Valley corn-fields (see p. 545 for an 
account of this). The feeding by ants on the fresh honey-dew can be readily 
observed in almost any garden (Fig. 247), and undoubtedly the mere presence 
in the aphid neighborhood of such redoubtable warriors as the ants is a 
strong deterrent of various predaceous insect enemies of the plant-lice. 
But most of the stories of ants and aphids printed in popular natural-history 
books need to be tested by careful observation. 

176 Bugs, Cicadas, Aphids, and Scale-insects 

Of all aphid species the grape-phylloxera, Phylloxera vastatrix (Fig. 248), 
has deservedly the widest notoriety. First made known in 1853 by Fitch from 
specimens found in New York, it was soon discovered to be well scattered on 
wild vines over the eastern United States. "It was introduced into the 
south of France before 1863, upon rooted vines sent from America; 
though the insect itself was not found and described there until 1868. 
The infection commenced at two points: one in the southeast in Card, 
the other in the southwest near Bordeaux. In 1868, when the nature of 
the pest was understood, it had already invaded considerable areas. The 

Fig. 248. — The grape-phylloxera. In upper left-hand corner an egg from which a male 
has issued, next an egg from which a female has issued; in upper right-hand corner 
winter egg; at left hand of middle row a just-hatched young, next a male (note 
absence of mouth-parts); at right end of middle row, female; lower figure, winged 
form. (After Ritter and Riibsaamen; much enlarged.) 

two areas first attacked gradually enlarged until they touched about the 
year 1880, and the insect began to spread northward. By 1884 about 
2,500,000 acres, more than one-third of all the vineyards of France, had 
been destroyed and nearly all the rest were more or less affected. The 
progress of the disease in parts of southern France was so rapid that in some 
towns vine-stumps became the principal fuel. Since 1884 the pest has 
continued to spread with somewhat less rapidity in France, partly because 
the most densely planted vineyard districts had already been devastated, 
but also because elsewhere its progress was retarded by quarantine and 
other restrictive measures. No remedies yet discovered, however, are 
capable of exterminating the pest; and to-day there is no vine-grow- 

Bugs, Cicadas, Aphids, and Scale-insects 177 

ing region of any importance in France, or elsewhere, exempt from 

Curiously enough this native American pest came to California, in which 
state it has done much more damage than elsewhere in our country, from 
France, introduced on imported cuttings or roots. It was tirst noticed about 
1874; by 1880 vines had been killed by phylloxera in three counties and 
hundreds of acres had been pulled up in the famous Sonoma Valley. 
Since then the pest has spread, according to Bioletti, to all the important 
grape-growing regions of central and northern California, and probably not 
less than 30,000 acres of vineyards have been destroyed. 

The phylloxera appears normally in four forms: (i) the gall form, living 
in little galls on the leaves, and capable of very rapid multiplication (this 
form rarely appears in California); (2) the root form, which is derived from 
individuals which migrate from the leaves to the roots, and which, by its 
piercing of the roots, sucking the sap, and producing little quickly de- 
caying tubercles on the rootlets, does the serious injury; (3) the winged 
form, which flies to new vines and vineyards and starts new colonies; and 
finally (4) the sexual forms, male and female, which are the regenerat- 
ing individuals, appearing after several agamic generations have been 

The life-history of the pest has been described as follows by Bioletti: 
"Some time during the summer, usually in July or August, some of the eggs 
laid by the root-insects develop into insects of slightly different form, called 
nymphs. They are somewhat larger than the normal root form and show 
slight protuberances on the sides, which finally develop into wings. These 
are the winged or colonizing insects, which emerge from the soil and, though 
possessing very weak powers of flight, are capable of sailing a short distance, 
and if a wind is blowing may be taken many rods or even miles. Those 
which reach a vine crawl to the under side of a leaf and deposit from three 
to six eggs. These eggs are of two sizes, the smaller of which produce males 
and the larger females. The female, after fertilization, migrates to the 
rough bark of the two-year-old wood, where she deposits a single egg, called 
the winter egg, which remains upon the vine until the following spring. 
The insect which hatches from this egg in the spring goes either to the young 
leaves and becomes a gall-maker, or descends to the roots and gives rise to 
a new generation of egg-laying root-feeders. The normal and complete 
life-cycle of the phylloxera appears then to be as follows: Male and female 
insects (one generation in autumn); gall-insects (one to five generations 
while the vines are in leaf); root-insects (an unknown number of genera- 
tions throughout the year); nymphs, which become winged insects (one 
generation in midsummer). The gall stage may be omitted, as it generally 
is in California, and the insects which hatch from the fertilized eggs laid by 

1 78 Bugs, Cicadas, Aphids, and Scale-insects 

the female go directly to the root and produce offspring which are in- 
distinguishable from the root form produced in the normal cycle. For 
how many generations the root form can exist and reproduce without the 
invigoration supposed to come from the production of the sexual form is 
not known, but certainly for four years and probably for more. The 

Fig. 249. — Roots and rootlets of grape-vine infested by the phylloxera. (After Ritter 
and Riibsaamen; enlarged.) 

gall form on American vines can be prevented by spraying the vines in 
winter with liquids to kill the winter eggs; but this treatment has no 
effect on the root forms, which in California hibernate abundantly in the 

All forms of the phylloxera species are very small, about -j'-g- of an inch 
being an average for fully developed individuals. The root form is light 
greenish yellow in summer, when it can be found by examining the rootlets 
of infested vines, and bronzy purplish in winter, when it can be found in 
little patches under the bark just at the crown of the vine. The newly 

Bugs, Cicadas, Aphids, and Scale-insects 179 

hatched young of the root form moves about freely, but when it reaches 
the egg-laying stage it becomes fixed. 

The chief injury to the vine is not sap-drinking, but the decaying or 
"cancer" of the roots caused by the punctures and tubercle forming (Fig. 
249). It usually takes tw^o or three years for phylloxera to kill a vine, but 
the results of the infestation are shown each season in the increasingly reduced 
growth of the new wood and in the lessened bearing. Suspected vines 
should be dug up and the rootlets carefully examined for tubercles and 
the insects themselves. The remedies, unfortunately, are either expensive, 
difficult, or severe. If a vineyard can be submerged for six weeks under 
at least six inches of water, the insects will be killed (by suffocation). Car- 
bon disulphide can be put into the soil among the roots by an injector at 
a cost of from ten to twenty dollars an acre. " This method succeeds only 
in rich, deep, loose soils and cannot be successfully used in soil containing 
much clay or on dry rocky hillsides." Finally, most severe but most effec- 
tive is the digging up of the whole of an infested vineyard and replanting 
resistant vines. "A resistant vine is one which is capable of keeping alive 
and growing even when phylloxera are living upon its roots. Its resistance 
depends on two facts: first, that the insects do not increase so rapidly on 
its roots; and second, that the swellings of diseased tissue caused by the 
punctures of the insects do not extend deeper than the bark of the rootlets 
and are sloughed off every year, leaving the roots as healthy as before. 
The wild vines of the Mississippi States have evolved in company with the 
phylloxera, and it is naturally among these that we find the most resistant 
forms. No vine is thoroughly resistant in the sense that phylloxera will not 
attack it at all; but on the most resistant the damage is so sHght as to be 
imperceptible. The European vine, Vitis vinifera L., is the most suscep- 
tible of all, and all the grapes cultivated in California, with a few unimportant 
exceptions, belong to this species." But the preferred French stocks can 
be grafted on to resistant American roots and the vineyard made practically 
immune. This is the method which has rehabilitated the French vine- 
yards and is now rehabilitating the -California ones. 

Another very important aphid pest of this country is the woolly 
apple-aphis, called in England and in Europe the American blight. This 
species, like the phylloxera, appears in different forms and lives both above 
ground on the twigs and larger branches and underground on the roots. 
It makes itself conspicuous and readily recognizable by the abundant fluffy 
waxen "wool" which it secretes. Badly attacked trees have the bark of 
their branches badly "cankered" and the roots covered with excrescences, 
and may die. The injuries are almost always severe, and the pest is one 
difficult to eradicate. If but few trees in an orchard are attacked, it is best 
to dig them up and burn them. The bark can be thoroughly sprayed or 

I 80 Bugs, Cicadas, Aphids, and Scale-insects 

scrubbed with a carbolized solution of soft soap (soap 10 parts, carbolic 
acid 2 parts, water 88 parts) and carbon disulphide injected into the soil about 
the base of the tree. 

Of the various aphids which attack foliage trees, the most familiar are 
those which resemble the woolly apple-aphis in their habit of secreting floc- 
culent masses of wax, and thus obtain the name of " blight," as elm-blight, 
beech-blight, alder-blight, etc. The alder-blight, or woolly alder-aphis. 
Pemphigus tessellata, gives birth in autumn to vast numbers which crawl 
down the trunks to the ground, where they congregate in the crevices between 
the base of the trunk and larger roots and the soil, or beneath the fallen leaves 
or other rubbish at the surface. They remain in their hiding-place until 
spring, when at the coming of the first warm days they crawl up the tree 
and out to the budding tips of the twigs. Here they begin sucking sap and 
at the same time secreting waxen "wool." In a week or so they become 
mature and begin giving birth to living young, and hereafter during the 
autumn and summer agamic generation after generation is produced. With 
the oncoming of cold weather the last generation crawls down to the ground 
to seek winter quarters. No sexual forms of this species have yet been 

Among the gall-forming aphids, one of special interest, because of the 
strange character and abundance of its galls, is the cockscomb gall-louse, 
Colopha ulmicola. Elm-trees infested by this aphis develop on the upper 
side of the leaves narrow, erect, blackish galls irregularly toothed along 
the top, and suggesting a cock's comb sufficiently to warrant the common 
name. These aphids secrete much honey-dew, noticeable on sidewalks 
under the trees and on the leaves, and in this honey-dew where it covers 
the galls and leaves grows a blackish fungus. 

Of all the families of the Hemiptera, probably the most important from 
the economic entomologist's point of view is that of the Coccidae, or scale- 
insects, and from the point of view of the biological student, also, no other 
is more interesting and suggestive. More nearly on a footing with the 
Coccids than any other Hemiptera are the Aphididas, just studied, but the 
scale-insects are even more specialized in curious and unusual ways, both 
as regards structure and physiology. In the more specialized scale-insects 
the females are quiescent in adult life, as well as in part of the immature 
life, and their fixed bodies are very degenerate, lacking both organs of loco- 
motion and of orientation, viz., eyes, antennae, wings, and legs. The family 
is a large and widely distributed one, numbering about 1450 known species 
in the world, of which 385 occur in the United States, but almost all are 
small and obscure and so foreign in appearance to the usual insect type 
that but few others than professional entomologists and the harassed fruit- 
growers ever recognize them as insects. Most of us have often had oppor- 

Bugs, Cicadas, Aphids, and Scale-insects 18 i 

tunily to make easy acquaintance with one or two species at our breakfast- 
tables; the t^attish, nearly circular little red-brown spots, or the more 
ovate blackish spots, which are occasionally to be seen on carelessly packed 
oranges are scale-insects and excellent examples of the extremie of degen- 
erate, quiescent type. The adult male scale-insects, unlike the females, 
are winged (although possessing but a single pair) and have eyes, an- 
tennae, and legs, but, strangely enough, no mouth-parts nor mouth-opening, 
so that they can take no food and must necessarily have but a few hours or 
perhaps days, at most, of life. And they are much more rarely seen 
than the females. Indeed, of many scale-insect species the males are not 
yet known, it being possible that in some species there is no male sex at 

The economic importance of the scale-insects has been keenly appre- 
ciated on the Pacific Coast ever since fruit-growing came to be a leading 
industry there, but the rest of the United States had not had to worry itself 
much because of the existence of these insect-scourges until recent years. 
A single Coccid species, however, has 
within ten years called the attention 
of entomologists and orchardmen 
and legislators all over the country to 
itself in a very illuminating manner. 
This species, the ill-named San Jose 
scale, Aspidiohis perniciosus, — which 
should rather be called "the perni- 
cious scale," or, if not that, then the 

Oriental scale, as it is a native of Japan 

^, . r . J 1 . Fig. 2t;o. — San Tose scale on bark of fruit- 

or China,— was first made known to ^^^^/ (After Slingerland; natural size.) 

science, and named, by Prof . J. H. Com- 

stock in 1880. Professor Comstock's specimens were collected in the Santa 
Clara Valley near San Jose, California. How much earlier the species 
had been brought to California is not known, but at the time of its naming 
by Professor Comstock it was already recognized by California fruit-growers 
as a serious pest, and Comstock wrote: "From what I have seen of it I think 
it is the most pernicious scale-insect known in this country." In August, 
1893, it was found to have got a footing in the east, and since then no other 
injurious insect — indeed hardly all others together — has received such con- 
stant and excited attention as has this obscure little pest. It is found now 
in every state and territory of the Union, and in Canada as well, and in 
thirty-five states has been the subject of hurried — and only partly w'ell- 
advised — legislation. This legislation has been directed toward restricting 
its spread by (a) quarantining it at the states' borders, and {h) inspecting 
orchards and nurseries for it within the state and attempting to stamp it 

I 82 Bugs, Cicadas, Aphids, and Scale-insects 

out. The structural characteristics and life-history of the insect may be 
briefly described as follows: 

There may be seen on infested branches, leaves, or fruit, small, flat, 
grayish, irregularly circular scales of varying size (Figs. 250 and 251), the 
large stones (about -^j inch diameter) being the adult females and the smaller 

ones being the immature individuals 
of both sexes. These circles are thin 
waxen plates, bearing one or more (de- 
pending on the age of the individual) 
faintly yellowish concentric inner cir- 
cles or plates (the inner one usually 
blackish and like a tiny nipple) which 
are the moulted exuviae of the scale. 
When the plant is badly infested the 
scales lie thickly together, even overlap- 
ping, and forming a sort of grayish 
scurf over the smooth bark. By rubbing 
or crushing this scurf a yellowish oily 
liquid issues from the injured bodies. 
If a scale be tipped over with a pin- 
point, there will be found underneath 
it a delicate flattened yellowish sac-like 
creature, the insect itself (Fig. 252). 
If adult, this degenerate female will be 
seen (by examination with magnifier) 
to have no distinct head, no eyes nor 
antennae, no wings nor legs. It does have a long, fine, flexible, thread-like 
process projecting from near the center of its under side; this is the suck- 
ing proboscis, and serves as a means of attachment to the plant as well 
as the organ of feeding. 

Early in the spring, females which have hibernated under their pro- 
tecting armor begin giving birth to living young, and continue doing this 
actively for about six weeks, when they die exhausted. The minute orange- 
yellow young, which have eyes, antennae, and three pairs of legs, crawl out 
from under the scale and run about actively for a few hours over the twigs 
or leaves; then they settle down and each* "slowly works its long bristle- 
like sucking-beak through the bark, folds its antennae and legs beneath its 
body and contracts to a nearly circular form. The development of the 
scale begins even before the larva becomes fixed. The secretion starts 

Fig. 251. — The San Jose scale, Aspi- 
diolus perniciosus, females and young, 
on bark of fruit-tree. (From living 
specimens; at left, natural size; at 
right, considerably enlarged.) 

* The following long quotation is made from Howard and Marlatt's " The San Jose 
Scale " (Bull. 3, N. S., Div. Ent., U. S. Dept. Agric, 1896). 

Bugs, cicadas, Aphids, and Scale-insects 183 

in the form of very minute white fibrous waxy filaments, which spring from 
all parts of the body and rapidly become more numerous and dense. At 
first the orange color of the larva shows through the thickening downy white 
envelope, but within two days the insect becomes entirely concealed by the 
white or pale grayish-yellow shell or scale, which now has a prominent central 
nipple, the younger ones often possessing instead a central tuft. The scale 
is formed by the slow matting and melting together of the filaments of wax. 
During the first day the scale appears like a very microscopic downy hemi- 
sphere. The matting of the secretion continues until the appearance of 
down and individual filaments is entirely lost and the surface becomes 
smooth. In the early history of the scale it maintains its pale whitish or 
grayish-yellow color, turning gradually darker gray, the central nipple 
remaining lighter colored, usually throughout development. 

"The male and female scales are exactly similar in size, color, and shape 
until after the first moult, which occurs twelve days after the emergence of 
the larva. With this moult, however, the insects beneath the scale lose all 
resemblance to each other. The males (Fig. 252, a) are rather larger than 
the females, and have large purple eyes, while the females have lost their 
eyes entirely. The legs and antennje have disappeared in both sexes. The 
males are elongate and pyriform, while the females are almost circular, 
amounting practically to a flattened sac with indistinct segmentation, 
and without organs, except a long sucking-bristle springing from near the 
center beneath. The color of both sexes is light lemon-yellow. The 
scales at this time have a decidedly grayish tint, overcast somewhat with 

"Eighteen days from birth the males change to the first pupal condition 
(propupa), and the male scales assume an elongate oval, sometimes shghtly 
curved shape, characteristic of the sex, the exuvia or cast larval skin show- 
ing near the anterior end. The male propupas are very pale yellow, with 
the legs and antennae (which have reappeared) together with the two or three 
terminal segments colorless. . . . Prominent wing-pads extend along the side 
of the body. 

"The female undergoes a second moult about twenty days from the 
larva. At each moult the old skin spHts around the edge of the body, the 
upper half adhering to the covering scale and the lower forming a sort of 
ventral scale next to the bark. This form of moulting is common to scales 
of this kind. 

"The covering scales at this stage are of a more purplish gray, the por- 
tion covering the exuviae inclining to yellowish. The male scales are more 
yellowish than the female. The effect of the sucking of the insects is now 
quite apparent on the young growth, causing the bark to assume a pur- 
plish hue for some distance around the central portion, contrasting strongly 

184 Bugs, Cicadas, Aphids, and Scale-insects 

Fig. 252. — The San 
Jose scale, Aspidiotiis 
perniciosus. a, male; 
/), a d u 1 1 female. 
(Much enlarged.) 

with the natural reddish green of the uninjured bark. With the second 
moult the females do not change materially from 
their former appearance, retaining the pale-yellow 
color with a number of transparent spots around 
the edge of the body. The sucking-bristles are 
extremely long, two or three times the length of the 
body of the insect. 

"About twenty days after birth the male insect 
transforms to the true pupa. With the first moult 
the shed larval skin is retained beneath the scale 
as in the case of the female; with the later moult- 
ings the shed skins are pushed out from beneath 
the scale. The scale, after the second moult, pre- 
sents on the inside two longitudinal ridges run- 
ning from one end to the other, touching the sides 
of the pupa, and which apparently enable the 
insect to move backward or forward and assist the imago in pushing itself 

"The true pupa is pale yellow, sometimes purplish, darkened about 
the base of the abdomen. The head, antennae, legs, wing-pads, and style 
are well formed, but almost colorless. . . . 

"From four to six days later, or from twenty-four to twenty-six days 
from birth, the males mature and back out from the rear end of their 
scales, having previously, for a day or two, remained practically developed, 
but resting under the scale. They seem to issue chiefly by night or in 
the evening. 

"The mature male (Fig. 252) appears as a delicate two-winged fly-like 
insect with long feelers and a single anal style projecting from the end of 
the body; orange in color, with a faintly dusky shade on the prothorax. 
The head is darker than the rest of the body, the eyes are dark purple, and 
the antennae, legs, and style are smoky. The wings are iridescent with 
yellow and green, very faintly clouded. 

"Thirty days from birth the females are full grown and the embryonic 
young may be seen within their bodies, each enclosed in a delicate mem- 
brane. At from thirty-three to forty days the larvae again begin to make 
their appearance. 

"The adult female, prior to the development of the young, measures 
one millimeter in length and a little less in breadth, and is pale yellow with 
transparent spots near the margin of the body (Fig. 252). 

"The length of a generation is determined by the female, and, as shown 
by the above record, covers a period of from thirty-three to forty days. Suc- 
cessive generations were followed carefully throughout the summer, and 

Bugs, Cicadas, Aphids, and Scale-insects 185 

it was found that at Washington four fuU generations are reguhirly developed, 
with the possibiUty of a partial fifth generation. On a number of potted 
trees a single overwintered female was left to each tree. After the full 
progeny of this individual had gone out over the tree all were removed 
again, except one of the oldest and fertilized females. This method was 
continued for each generation throughout the breeding season. Some 
interesting records . . . were thus obtained, which indicate the fecundity 
of the females as well as the number of generations." 

From these records it may be fairly estimated that an average of 200 
females (in addition to about as many males) are produced by each female, 
and that there are four generations each year in the latitude of Washington, 
D. C. Thus the product of a single overwintered female in a single year 
amounts to 3,216,080,400 male and female descendants. This total is, 
of course, never reached, because only a part of each generation reaches 
maturity and produces young, but in a favorable season on a tree newly 
infested (and thus providing a plentiful food-supply) a large majority of 
each generation do most probably go through their normal existence. 
"Neither the rapidity with which trees become infested," add Howard and 
Marlatt, "nor the fatal effect which so early follows the appearance of this 
scale-insect is therefore to be wondered at." 

But not all scale-insects are so specialized either structurally or physio- 
logically as the pernicious (or San Jose) scale. The females of some species 
retain the eyes, antennae, and legs through their whole life and can crawl 
about if need be at any time. Others show a sort of transition between 
these two extremes of activity and quiescence, having the legs present, but 
in adult life much reduced in size and probably functionless, or at best 
capable of carrying the insect but feebly and briefly. In the matter of the 
covering, too, there is much variety; some scales secrete no wax at all, but 
have the body-wall of the back specially thickened and made firm so as 
to act as an effective covering-shield underneath which, somewhat as with 
a turtle, the legs and head can be concealed. Others secrete filaments or 
tufts of soft white wax which form a sort of felted protecting covering for the 
body. In a general way the various scale-insects may be instructively 
gathered into three groups, depending on the characters of the females; 
in the first group the females retain the antennae, eyes, and legs, and the 
segmented condition of the body (typical of normal insects) and are capable 
of locomotion throughout life; they secrete wax usually in the shape of white 
cottony filaments or masses with which they cover the body more or less 
completely, sometimes forming conspicuous waxen egg-sacs at the posterior 
extremity of the body; the females of the second group retain their legs 
and antennae through life, but have them in reduced condition when adult, 
and although capable of feeble motion, usually lie quiescent; they commonly 

1 86 Bugs, Cicadas, Aphids, and Scale-insects 

Fig. 253. — The fluted or cottony 
cushion-scale, Icerya purchasi, 
winged male and wingless 
female with fluted waxen egg- 
sac {es). (After Jordan and 
Kellogg, much enlarged.) 

secrete no wax, but have the body-wall of the dorsum strongly chitinized, 

and usually very convex, so that it forms 
a strong rigid protecting shell; finally the 
females of the third (and largest) group are 
the so-called armored scales, which in the 
adult stage are degenerate creatures without 
distinct body segmentation, without antennae, 
eyes, and legs, thus being incapable of 
locomotion; they form a flatfish or convex 
dorsal scale of secreted wax and of the cast 
skins or exuviae of the body. 

In all the groups the males (Figs. 252 and 
253) are very different in appearance from the 
females, being minute fly-like creatures with 
a single pair of wings, a pair of long antennae, 
and a plump, soft, little body, usually 
terminating in a single needle-like process or 
in a pair of long waxen hairs. Males are 
not yet known for some of the species. 
Familiar examples of the first group are the mealy-bugs {Dadylopiits sp.) 
of greenhouses and gardens, soft-bodied scales, bearing projecting rods 
and threads of white wax of varying length, and rather prettily arranged. 
A more famous and interesting member of this group is the fluted or cottony 
cushion-scale, Icerya purchasi (Fig. 253) (so called because of the beautiful 
fluted white waxen egg-sac secreted by the female), which once threatened 
to destroy all the orange-groves of California, but was brought to bay by 
a little red and black ladybird-beetle, Vedalia cardinaJis (Fig. 254), brought 
from Australia for this very purpose. In 1868 some young orange-trees 
were brought to Menlo Park (near San Francisco) from Australia. These 
trees were undoubtedly infested by the fluted scale, which is a native of 
Australia. These scale immigrants throve in the balmy California climate, 
and particularly well, probably, because they had left all their native enemies 
far behind. By 1880 they had spread to the great orange-growing districts of 
southern California, five hundred miles away, and in the next ten years 
caused enormous loss to the growers. In 1888 the entomologist Kcebele, 
recommended by the government division of entomology, was sent at the 
expense of the California fruit-growers to Australia to try to find and send 
back some effective predaceous or parasitic enemy of the pest. As a result 
of this effort, a few Vedalias were sent to California, where they were zeal- 
ously fed and cared for, and soon, after a few generations, enough of the little 
beetles were on hand to warrant trying to colonize them in the attacked 
orange-groves. With astonishing and gratifying success the Vedalia in a 

Bugs, Cicadas, Aphids, and Scale-insects 187 

very few years had so naturally increased and spread that the ruthless scale 
was definitely checked in its destruction, and from that time to this has 
been able to do only occasionally and in limited locaHties any injury at all. 

Fig. 254.— The fluted scale, Tcerya pitrchasi, attacked by the Australian ladybird-beetle, 
Vedalia cardinalis. In lower left-hand corner a Vedalia which has just issued from 
its pupal case. (From life; upper figure slightly enlarged; lower figiire much 

Of the second group, the best-known scales are the various species of the 
genus Lecanium (Fig. 256). Of these, the olive or oleander or black scale, 
L. olece, as it is variously called, is the most widely distributed and abundant 
and hence economically important. It is a long-known species, having 
been described in Europe in 1743, and it was brought to this country in 
early days. The adult females are blackish, almost hemispherical, rough- 
skinned creatures, with no external indication of head or other body divi- 
sions, feet, antennae, etc., all these parts being visible only from the ventral 
aspect, which normally is closely applied to the leaf or twig. On the back 
may be distinguished three ridges forming an H. The young are flatter 
and light brown, but can be recognized by their even more distinct H-mark. 
This scale is found all over the United States and has a wide range of food- 
plants, garden-bushes of all kinds, as well as deciduous and citrus fruits 
being attacked. In California it is one of the worst insect-pests of the olive- 
tree and also one of the worst of the orange enemies. It has certain natural 
enemies in the persons of various ladybird-beetle species, and a few special 
ladybird-beetles have been imported from Australia and elsewhere in the 
hope of repeating the signal Vedalia success. Only a fair measure of suc- 
cess has been achieved. An indirect but serious injury caused to plants 
by the black scale is due to the germination in the honey-dew secreted by 
it of the spores of a fungus, Capnodium sp., which spreads its felted mycelia 

J 88 Bugs, Cicadas, Aphids, and Scale-insects 

over the leaf-surfaces, closing the breathing-pores (stomata) and thus truly 
suffocating the plant. Although this scale species has been known for a 

century and a half, the males have 
been seen but few times and in but 
few places. Another familiar member 
of this group, which secretes a distinct 
white waxen egg-sac, is the maple- 
scale, Pith'inaria inmimerahihs (Fig. 
255), common on maples in the 
eastern states. 

Of the third group, that of the 
most specialized (degenerate) scales, 
the pernicious scale, already fully 
described, may be taken as a shining 
example. There is a host of these 
armored scale-insects, and few trees or 

Fig. 255. 

Fig. 256. 

Fig. 255. — The maple-scale, Pulvinaria innumerahilis. Females with egg-sacs on the 
twig; young scales on under side of leaf, and a single young scale, much enlarged, 
at left. (After Felt; natural size.) 

Fig. 256. — Lecaniiim scales attacked by the fungus Cordyceps clavidata. (After Pettit; 
much enlarged.) 

shrubs escape their attacks. The various genera are mostly distinguishable 
by the shape of the covering scale, but to determine the species exactly 
requires, for many, careful examination, under high powers of the microscope, 
of the minute chitinous processes which form a fine fringe along the posterior 
margin of the last abdominal segment. To make this examination it is 
necessary to remove the female from under her scale, and mount her cleared 
body flat in balsam or glycerine on a glass slide. An important species 
in this group is the red orange-scale, Aspidiotus aiirantii (Fig. 257), common 
in orange-groves of southern California. A species very closely resembling 
it is A. ficus, common in the Florida groves. On pine-needles one may 
often note small, narrow elongate white waxen scales, with the smaller, 
yellowish-brown exuviae at one end; these belong to the widely spread species 
Chionaspis pinifolice. On apple-trees often occurs a roughened shining 

Bugs, Cicadas, Aphids, and Scale-insects 189 

Fig. 257. — The red orange-scale, Aspidiotus 
aiirantii. a, females, natural size, on leaf; b, 
female, much enlarged, removed from under 
waxen scale; r, the scale, composed of wax 
and exuviae, much enlarged; d, just hatched 
young, much enlarged; e, male, much enlarged. 
(After Jordan and Kellogg.) 

blackish narrow elongate curved scale, resembling a little an oyster-shell 

in miniature; this is the sometimes serious apple-pest, Mytilaspis poniorum. 

But we have no space to list 

even the most important of 

these degenerate but successful 

insect enemies of our fruit- and 


The devising of remedies for 
scale attack has been given much 
attention, and a number of effec- 
tive means have been discovered 
for fighting the pests. Probably 
the most effective of all is the 
fumigation of infested orchard- 
trees by hydrocyanic gas. A 
tent capable of enclosing a whole 
tree is made, and with this in 
place hydrocyanic gas is gen- 
erated under it by pouring 
about 50 oz. of water into 5 oz. 
of commercial sulphuric acid and 
dropping in 15 oz. of cyanide of potassium, these amounts of acid, water, and 
cyanide being sufficient to fumigate a tree 12 ft. high by 10 ft. in foliage diam- 
eter; that is, to fumigate about 1000 cu. ft. of space. For larger or smaller trees 
change the amounts of acid, water, and cyanide proportionally. Of washes 
to be applied in winter, when the leaves are off, the best is one made of lime 
50 lbs., sulphur 50 lbs., salt 50 lbs., water 150 gals.; slake the lime with 
water enough to do it thoroughly, and during the process add the sulphur. 
Boil one hour with water enough to prevent burning and until the mixture 
becomes of a deep amber color. Dissolve the salt in water enough to do 
it quickly and add slowly to the boiling mass. When all is thoroughly 
mixed together and has actually boiled at least an hour add water enough 
to make up 150 gals., and apply by spraying or washing while hot. It 
may be safely applied when the foliage is off to any fruit-tree, garden shrub, 
or small fruit, and is a very effective "scale-killer." Of sprays for the leaves, 
crude petroleum and kerosene emulsion are the best. For use, undiluted 
crude petroleum should be entirely untreated and of specific gravity of 43° 
or over on the Beaume scale. Smith has used this oil safely on all ordinary 
fruit-trees, but advises not applying it to peach-trees. At time of apply- 
ing, the trees' should be dry, the oil of a temperature not below 60° Fahrenheit, 
and the nozzles should throw a perpetual fine spray. Kerosene emulsion is 
made by boiling J lb. of hard soap in i gal. of water and then adding 2 gals. 

190 Bugs, Cicadas, Aphids, and Scale-insects 

Fig. 258. — Female red orange-scale, Aspi- 
diotus aiirantii. (Photomicrograph by 
George O.Mitchell; much enlarged.) 

of kerosene and churning violently until a thick white cream is formed. 
Let this cool and jelly; it is the "stock," and will hold for a few weeks; when 

ready to spray, dilute stock with twelve 
to fifteen times its own bulk of water 
and spray finely over the foliage. The 
spraying should be done when the 
young scales are hatching and crawl- 
ing about. They are then easily killed 
by contact with even a single fine drop 
of kerosene. For peach-trees dilute 
the stock twenty times. 

Some of the scale-insects present 
such unusual conditions of structural 
modification and of habits that they 
are, when first met with, difficult to 
recognize as insects at all. The 
waxen covering may be so irregular and 
curiously shaped that it gives no clue to the character of the enclosed insect 
(Fig. 261), but seems to be simply a secretion of the plant in which the insects 
are found. Or the globular shape and absence of distinct body-parts may 
make the insects with their hardened blackish cuticle look like small plant- 
galls; indeed certain scale-insect species were first described by botanists as 
galls. Some scales live under ground, either in 
ants' nests or independently; the curious so-called 
"ground-pearls," small spherical shining bodies 
found loosely scattered in the soil in certain tropic 
regions, and really collected to be strung on threads 
or necklaces, are the strangely modified bodies of 
Margarodes jormicarum, a scale-insect. Taken alto- 
gether, probably no other family of insects exceeds 
the Coccidae in the extremes of strange specializa- 

Closely related to the plant -lice and scale- 
insects are the mealy- winged flies, constituting the 
family Aleyrodidae. The adults (Fig. 262), except 
of two or three of the most abundant species, are 
rarely seen even by professional entomologists, but 
careful search will reveal in almost any locality the 
curious little box-like elliptical bodies of the young 
(Fig. 263), usually shining black, with pure-white 
waxen rods, filaments, or tufts. Examined under a good magnifier, the 
wax-tufted cases are exquisite objects. These young mealy-wing flies look 


Fig. 259. — Female rose- 
scale, Diaspis r o see. 
(Photomicrograph b y 
George O. Mitchell; 
much enlarged.) 

Bugs, Cicadas, Aphids, and Scale-insects 191 

much like scale-insects and have the same general habits. Provided with 
a delicate long sucking-beak, each individual remains fixed in one spot on 
a green leaf, sucking up its food, the plant-sap, as it needs it. The adults 
which finally issue from the beautiful little cases have four rounded wings, 
pure white or with small dusky spots and golden yellow, finely beaded 
margins; each wing has but a single vein, and is dusted with a granular 

Tig. 260. — The California live-oak scale, Ccrococcus ehrhonii. 

Patterson; natural size.) 

(,l'huK)graph by Rose 

white waxen powder or "bloom." The tiny white or pale-yellow eggs 
are laid on leaves in a circle or the arc of one, in one or more rows, and 
vary in number from three to thirty; each egg has a minute but noticeable 
curving stem. The young hatch in from ten to thirteen days, and move 
freely about, but never seem to get more than about one inch from the 
deserted shells. This activity lasts for from ten to forty hours; then 
the young attach themselves to the leaf by inserting the sucking proboscis, 

192 Bugs, Cicadas, Aphids, and Scale-insects 

and soon moult, losing at this time the legs and antenna. After a second 
moulting, however, minute new legs and antennas are again to be seen, and 
later the wing-pads appear, and wings, legs, and antennae develop and grow 
apace; at a last moulting the insect leaves the protection of its beautiful little 
case and flies away. Leaving the pupa-case is a slow and toilsome process, 
the imago often struggling for hours before it is free and ready for flight. 


261. — The Southern California oak-scale, Cerococcus qiiercus. 
(Photograph by Rose Patterson; natural size.) 

All of the pupas secrete "honey-dew," sometimes in such quantities that 
the leaf around the case, and the top of the case itself, are covered with it. 
This honey-dew is emitted from the tip of a little flap-like anal structure called 
the lingula (Fig. 266). The sweet liquid honey-dew, when exposed to the 
air, becomes thick and finally hardens. The spores of fungi often germinate 
in the excreted honey-dew, and numerous ant-species collect it for food. 

Bugs, Cicadas, Aphids, and Scale-insects 193 

To distinguish any of the various species of mealy-winged flies would 
be a difficult matter for the beginning 
entomologist. Two special students of 



Fig. 262. 

Fig. 263. 

Fig. 262. — A mealy-wing, Aleyrodes pruinosa, adult. (After Bemis; much enlarged.) 
Fig. 263. — Pupa of Aleyrodes tentacidatus. (After Bemis; much enlarged.) 

Fig. 264. Fig. 266. 

Fig. 264. — Pupa of Aleyrodes iridescens. (After Bemis; much enlarged.) 
Fig. 265. — Pupa-case of Aleyrodes merlini. (After Bemis; much enlarged.) 
Fig. 266. — Vasiform orifice and Hngula of pupa of Aleyrodes merlini. (After Bemis; much 

the American species have published lists and descriptions of all the kinds 
so far known in this country, namely, Quaintance (Bull. 8, Tech. Ser., Div. 
of Ent., U. S. Dept. Agr., 1900), who has studied the eastern species, and 

194 Bugs, Cicadas, Aphids, and Scale-insects 

Bemis (Proc. U. S. Nat. Mus., vol. 27, 1904), who has studied the Pacific 
Coast forms. Mrs. Bemis found twenty hitherto unknown species of mealy- 
winged flies in easy collecting 
range of Stanford University, 
and these twenty kinds added 
to those already known make a 
total of sixty different species so 
far recorded from the United 
States. There are certainly 
many more species yet unde- 

The mealy-winged flies have 
some, though not a large, eco- 
nomic importance. One or two 
species, Aleyrodes vaporariorum, 
Fig. 267.— Pupa of Aleyrodes merlini, showing etc., are recognized as pests in 
long waxen tufts. (After Bemis; much en- greenhouses; one, A. citri, is a 
larged.) - , , 

pest of oranges, and another, 

A. packardi, injures strawberry-plants. In all these cases probably as much 
injury is done by the suffocating fungus growth that is supported by the 
secreted honey-dew as by the direct sap-sucking of the Aleyrodes themselves. 
Fumigation by hydrocyanic gas (see p. 189) is probably the best remedy 
for the greenhouse and orange mealy-wings, and spraying with kerosene 
emulsion (see p. 189) the best for the strawberry Aleyrodes. 


Key to Families of the Heteroptera (includes both Nymphs and Adults). 
(Adapted from Woodworth, with some Additions.) 

Antcnnse shorter than the head: aquatic or shore insects. 

With two ocelli (Toad-bugs.) Galgulid^.' 

With no ocelli. 

Hind feet without claws; aquatic insects. 

Prothorax overlapping the head above (Back-swimmers.) Notonectid^. 

Head overlapping prothorax above (Water-boatmen.) Corisid^. 

Hind feet with claws. 

With two long processes on tip of abdomen which can be held together to form 

a tube (Water-scorpions.) Nepid^. 

Without abdominal processes, or if any, short flattish retractile ones. 

Hind legs broad and flat (Giant water-bugs.) Belostomatid^. 

Hind legs slender Naucorid^.'^ 

Antennae at least as long as the head: a few aquatic forms, but mostly terrestrial. 
Head as long as the whole thorax (Marsh-treaders.) Limnobatid^.'^ 

Bugs, Cicadas, Aphids, and Scale-insects 195 

Head shorter than thorax. 

Last segment of foot divided and the claws not at the tip. 

Middle and hind legs very long (Water-striders.) Hydrobatid^. 

Middle and hind legs not very long Veliid^. 

Last segment of foot not divided, and the claws at the tip. 
Antennae 3- or 4-segmented. 

Proboscis (or beak) with three joints. 

Body very long and slender (Thread-legged bugs.) Emesid.e. 

Body not long and slender. 

Femora of fore legs very wide (.\mbush-bugs.) Phymatid.-e. 

Femora of fore legs not very wide. ^ 

Antennas of three segments (Assassin-bugs.) Reduviid^. 

Antennae of four segments. 

Feet of two joints, and body very flat (Flatbugs.) Aradid^. v 

Feet of three joints. 

Body flat on top (Bedbugs.) Acanthiid.e. ' 

Body not flat on top (Shore-bugs.) Saldid.e. 

Proboscis (or beak) with four joints. 
Without ocelli. 

Heavy-bodied insects, membrane of wings (in adults) with two large cells 
at the base from which arise about eight branching veins (Fig. 268, 2). 

(Redbugs.) Pyrrochorid^. "^ 
Light-bodied insects; membrane of wings (in adults) with one or two closed 
cells at the base and with no longitudinal veins (Fig. 268, i). 

(Leaf- and flower-bugs.) Capsid.e. '' 
No ocelli. 

Fore legs very different from the others; wings when present in fully de- 
veloped condition with four long veins in the membrane bounding three 
discal cells, which are often open; from these cells diverge veins which 
form several marginal cells (Fig. 268, 5). . - (Damsel-bugs.) Nabid^. 
Fore legs not very different from the others. 

Body very narrow (.Stilt-bugs.) Berytid.e. 

Body not very slender. 

Feet of two joints; wing-covers (of adults) resembling lace network. 

(Lace-bugs.) Tingitid.e. 
Feet of three joints. 

Antennae inserted below an imaginary line drawn from the eye to the 
beak ; membrane of wing (in adults) with four or five simple veins 
arising from its base, the two inner veins sometimes joined to a 
cell near the base (Fig. 268, ,5) . . (Chinch-bug family.) Lyg.5:id.e. 
Antennae inserted above an imaginary line drawn from the eye to the 
beak; membrane of wings (in adults) with many usually forked 
veins, springing from a transverse basal vein (Fig. 268, 4). 

(Squash-bug family.) C0REID.E. 
Antennae 5-segmented. 
Body flat above. 

With few or no spines on the tibias (Stink-bugs.) Pentatomid.«. 

With rows of spines on the tibiae (Burrower-bugs.) Cydnid.e." 

Body strongly convex above. 

Prothorax round in front and nearly straight behind. 

(Negro-bugs.) Corimel^nid^. ' 
Prothorax hexagonal (Shield-backed bugs.) Scutellerid.t.. 

196 Bugs, Cicadas, Aphids, and Scale-insects 

We come now to the "true bugs," representing twenty-six families and 
constituting the Heteroptera, the largest of the three suborders of the 
Hemiptera. The classification of the members of this large group into 
families, by the use of the keys commonly used by entomologists, demands 
the recognition of such small and obscure structural characters that I have 
tried to find some easier means for the use of the amateur and general col- 
lector. As collecting and observing in the field imply the discovery of 
insects in their native haunts, we may acceptably make use of constant 
habits for a basis of convenient grouping. About one-third of the Heterop- 
terous families are aquatic in habitat, and of these the members of some 
are to be found on the surface of pools and ponds, of others swimming 

Fig. 268. — Wings of Heteroptera, showing disposition of veins in membrane character- 
istic of various families: i, Capsidae; 2, Pyrrhocoridae; 3, Lygasidas; 4, Coreida: 
5, Nabidae; 6, Acanthiidae. (After Comstock.) 

or crawling about below the surface, and of two, only partly aquatic, 
on the shore, but always by the water's edge. Some of these aquatic bugs 
are to be discovered occasionally in flight far from water, as the giant 
water-bugs and others, when circling about electric lights or in search of 
new homes. But the structural signs of the aquatic habitat, legs flattened 
and fringed so as to be fitted for swimming, will betray these estrays. 
Occasionally, too, a strictly terrestrial bug will be found on the surface of 
a pool, but his violent and obviously unaccustomed and awkward attempts 
to swim to shore will betray him. So we may begin an acquaintance with 
the Heteroptera by resorting to the nearest pond or quiet stream-pool. 

On the surface are the familiar water-striders, or skaters. Their long, 
spider-like legs, narrow and black or oval and yellow and black body, 
and swift nervous running distinguish them from all other bugs. They 
are members of the family Hydrobatidse, and the commoner species belong 
to the genus Hygrotrechus (Figs. 269 and 270). Upheld by the tense surface- 
film of the water, their feet only make little dimpled depressions in the sur- 
face, the shadows of which may often be seen on the sandy bottom. The 
locomotion is really due to a sort of surface rowing or gliding, and not a 

Bugs, Cicadas, Aphids, and Scale-insects 197 

true running. The water-striders are predaceous, capturing smaller living 
insects by running or leaping, and, with the prey held securely in the grasp- 

FiG. 269. Fig. 270. 

Fig. 269. — Water-strider, Hygrotrechus sp., adult. (Twice natural size.) 
Fig. 270. — Water-strider, Hygrotrechus sp., young. (Twice natural size.) 

ing fore legs, piercing and sucking the blood of the unfortunate victim, yet 
alive. Care should be taken in handling water-striders, as the sharp beak 

Fig. 271. Fig. 272. Fig. 273. 

Fig. 271.— Broad-bodied water-strider, Stephania picta. (After Uhler ; natural 

Fig. 272. — An ocean water-skater, Halobates wiil/ersdotffi, from near Galapagos Islands. 

(Three times natural size.) 
Fig. 273. — A marsh-treader, Limnobates lineata. (One and one-half times natural size.) 

can make a painful puncture. Some of them are winged and some wing- 
less, and both kinds of individuals may belong to the same species. The 

198 Bugs, Cicadas, Aphids, and Scale-insects 

young are usually short-bodied, and of course wholly wingless or with small 
wing-pads only. In late autumn the water-striders conceal themselves 
in the mud beneath leaves or rubbish or at the bottom of the pool under 
roots or stones to hibernate, coming out again with the first warm days of 
spring. The whitish elongate eggs are laid in early spring, being attached 
by a sort of glue to the leaves and stems of aquatic plants. Some species 
have several generations each year. Water-striders are easily kept in 
aquaria if the sides are high enough above water to prevent their leaping 
out. In bringing them in from the pond covered pails should be used, or 
they may be enclosed in any small dry receptacle not air-tight. They are 
easily drowned if shaken about in a covered pail of water. 

A few interesting Hydrobatids, belonging to the genus Halobates (Fig. 
272), live on the surface of the ocean, especially in subtropic and tropic 
latitudes. They are said to feed on the juices of dead animals floating on 
the surface, and probably attach their eggs to floating seaweed (Sargassum). 

Certain stout-bodied insects, widest across the prothorax and with much 
shorter, stouter legs, members of the family Veliidae, are sometimes to be 
found, running about on the surface of the water, always near the shore. 
They can also run readily on land, which the true water-skaters cannot 
do. Also certain other slender insects, about ^ inch long, with thin long 
legs and hair-like antennae and long cylindrical head, are to be found on 
top of the water. But they creep slowly about on the surface or on the 
soft mud of the shore, and are found mostly where plants are growing in 
quiet water. These are marsh-treaders, Limnobates lineata (Fig. 273), 
and this species is the only representative of the family Limnobatidae known 
in this country. 

Swimming and diving about beneath the surface are the water-boatmen 
(family Corisidas) and back-swimmers (family Notonectidae). The water- 
boatmen (Fig. 274) are oval, finely mottled, greenish gray and black, and 
swim with back uppermost. They are all small, some only \ inch long, 
none over half an inch. The back-swimmers have the back shaped like the 
bottom of a boat, swim with the back always down, and are usually bluish 
black and creamy white in color. Both of these kinds of water-bugs are 
predaceous, feeding on smaller aquatic creatures. But the beak of the back- 
swimmers is much longer and stronger than that of the water-boatmen, 
and can make a painful sting on one's finger. Both kinds have the hind 
legs long and specially flattened and fringed to serve as oars, and both kinds 
come to the surface for air, although the back-swimmers come up far more 
often than the water-boatmen. The air taken up clings as a silvery bubble 
to a large part of the body both under the folded wings and on the under 
side, being held there by fine hairs which form a pile like that on velvet. 
A supply of air is thus taken down by the bugs, which enables them to remain 

Bugs, Cicadas, Aphids, and Scale-insects 199 

for some time under water. Both kinds are attracted to lights, and may 
often be seen in summer about outdoor electric lamps. The eggs of the 
water-boatmen are attached to the submerged stems of aquatic plants, while 
those of the back-swimmers are inserted in the stems, the female having 
a sharp ovipositor for this purpose. In winter the adults lay dormant in the 
mud at the bottom of ponds or streams. 

All the species of water-boatmen in the country belong to the genus 
Corisa, while there are three genera of back-swimmers, Notonecta, with 
hind legs longer than the others and fore wings but little longer than the 
abdomen, being the most abundant and 
wide-spread. Plea is a genus with all the 
legs ahke, while Anisops, the third genus, 
has the wing-covers usually much longer 
than the abdomen. The complete life- 
history of no member of either of these 
families of water-bugs is yet known, but it 
ought not to be a difficult matter for some 
persistent observer to add this needed ^^^ ^^^ _^ wat^oatman, Corisa 
knowledge to entomological science. Both sp. (After Jenkins and Kellogg; 
water-boatmen and back-swimmers live twice natural size.) 
readily in aquaria, and make thoroughly interesting creatures to observe 
at leisure. The characteristic habits of obtaining air, swimming, capturing 
prey, etc., can all be learned from the observation of aquarium specimens. 
The capacity of the water-boatmen to remain below the surface in pure 
water for protracted periods, apparently indefinitely long, needs to be better 
understood than it is at present, and should be an interesting problem for 
some observer of aquarium life. 

Creeping or crawling about among the stems and leaves of submerged 
plants in reedy and grassy quiet waters, and feeding on smaller insects, may 
sometimes be found certain small flat-bodied oval insects with front legs 
thickened and fitted for grasping. These are water-creepers, or Naucoridae, 
only five species of which are known in this country. The single species 
found in the eastern states is known as Pelocaris jemorata, and is about 
J inch long, broadly oval in shape, and yellowish brown in color. The 
other species belong to the genus Ambrysus and are restricted to the western 
states. The life-history of no member of the family is known. 

Occasionally there will be seen resting, or swimming slowly about, at the 
bottom of the pool a veritable giant bug, 2f inches long and i J inches wide, 
with heavy strong legs flattened and oar-like and the front ones held out 
arm-like and bent in an expectant grasping position. Again, in the warm 
sultry evenings of midsummer and early autumn, among the swarms of 
insects attracted to the electric lights on the streets, one or two great bugs 

200 Bugs, Cicadas, Aphids, and Scale-insects 

will go whirling around the bright globe of light, casting large fleeting 
shadows on the ground below. The giant in the pool's depth and the giant 
in the giddy swarm at the light are one and the same, viz., the giant water- 
bug or electric-light bug, a member of the family Belostomatidae. Most 
of its life is passed in the water; it hatches from eggs deposited under water, 
lives its whole immature life in the pool, and only comes out for a short flying 
season to find mates or a new pool. The two largest species of this family, 
both common in this country, are Belostoma americana (Fig. 275) and Benacus 

griseus, distinguishable by the fact that 
the former has a groove on each front 
femur for the tibia to fit in when folded. 
A smaller kind, more oval in shape, is the 
commonest form on the Pacific slope. 
This is Serphus diJatatus, the toe-biter. 

Fig. 275. 

Fig. 275. — The giant water-bug or electric-light bug, Belostoma americana. (Natural 

Fig. 276. — The western water-bug, Serphus sp.; male with eggs deposited on its back 

by female. (Natural size.) 

which is i\ to j\ inches long and f to |^ inch wide. In the East a still 
smaller form, Zaitha fluminea, is common. This is a little less than i 
inch long. All these Belostomatids are fiercely predaceous, capturing 
aquatic insects, tadpoles, etc., and are armed with a short, strong, pointed 
beak with which a serious puncture can be made. They secrete themselves 
beneath stones or rubbish, whence they dart out on their victims. A con- 
siderable amount of poisonous saliva enters the wound made by the beak, 
and probably aids in overcoming the prey. The larger species attack 
young fish, seizing them with their strong grasping fore legs and sucking 
their blood. They can do much injury in carp-ponds or in garden-pools 
where fishes are kept for pleasure. The females of the species of the 

Bugs, Cicadas, Aphids, and Scale-insects 201 

smaller genera Serplnis and Zaitha have the curious habit of gluing their 
eggs upright, in a single layer, on the back of the unwilling male (Fig. 276). 
For a long time it was believed, and is so stated in most entomological books, 
that the female deposited the eggs on her own back, but it was discovered 
by Snodgrass that the female Serphits had no ovipositor capable of reach- 
ing to her back, and by Miss Slater that the female Zaitha is in similar con- 
dition. Miss Slater observed the egg-laying by aquarium specimens. The 
male struggles against the indignity, but is actually overcome by the female. 
Another small aquatic family of few species is that .of the Nepidae, or 
water-scorpions. These dirty brown, stick-like insects can be distinguished 
from other aquatic Hemiptera by the long slender respiratory tube, made 

up of separable halves each grooved on its 
inner face, which projects from the tip of 
the abdomen. Rather sluggish in habit, 
they lie at the bottom of a shallow pool and 
lift this respiratory tube up so that its open 
tip reaches the surface. They are preda- 
ceous and have the fore legs modified for 
seizing prey, the other legs being fitted for 
walking or crawling over the bottom. There 
are two common genera in the family: Nepa, 
with flattened oval body less than three times 
as long (not including respiratory tube) as 

Fig. 277. 

Fig. 278. 

Fig. 277. — A water-scorpion, Ranatra jiisca. (One and one-half times natural size.) 
Fig. 278. — Eggs of the water-scorpion, Ranatra jusca. (After Pettit; enlarged.) 

broad, and Ranatra (Fig. 277), with elongate slender body more than five 
times as long as broad. Like the giant water-bugs the water-scorpions 
lie in wait for their prey, trusting to their inconspicuous color and partial 
concealment in the mud and rubbish of the bottom to hide them from 
approaching victims. 

20 2 Bags, Cicadas, Aphids, and Scale-insects 

By the edge of pond or stream may be found representatives of two other 
small families, most striking in appearance and manner, the dark-colored, 
squat, broad, rough bodied, big-eyed, leaping toad-bugs (Galgulidae) and 
the smaller, soft, long-oval, long-legged, running shore-bugs (Saldidae). 
One species of toad-bug, Galgidiis ocidatiis (Figs. 279 and 280), is common 
all over the country and may often be found in considerable nimibers on 
the muddy banks of streams and ponds. It lives 
upon other insects, which it catches by creeping 
slowly to within a short distance and then suddenly 
leaping upon and seizing them with its strong front 


Fig. 279. 

Fig. 280. 

Fig. 279. — The toad-bug, Galgulus oculatus. (Three times natural size.) 
Fig. 280. — Three toad-bugs, Galgulus oculatus, "coming on." (From life; three times 
(natural size.) 

legs. Toad-bugs vary in general coloration with the mud or soil they are 
on, so as to harmonize with the ground color and thus be undistinguishable. 
The shores of a small pond, Lagunita, on the campus of Stanford 
University, vary much in ground color, three shades, namely, reddish, slaty 
bluish, and mottled sand color, being the principal 
ones, and toad-bugs collected from the banks of 
this pond show very noticeably all these distinct 
schemes of color. The shore-bugs (Saldidae) are 
represented by but one genus, Salda (Fig. 281), of 
thirty or more species, in our country. The insects are 
about y\- inch long, smooth-bodied, and narrower than 
the toad-bugs, blackish with white or yellow markings, 
and have long slender antennae. They prefer stream 
or pond banks which are weedy or grassy and offer 
They are common also on seabeaches. They feed 
on drowned flies and other insects, from which they suck the blood. They 
thus do some good as scavengers. 

The preceding ten families include all of the aquatic and strictly shore- 
inhabiting Hemiptera. The remaining sixteen families of the suborder 
Heteroptera, as well as all the families in both other suborders, are terres- 
trial, being found for the most part (the Parasita wholly excepted) on vegeta- 
tion where food, either the juices of the plants, or the blood of other plant- 

FiG. 281. — A shore- 
bug, Salda sp. (Six 
times natural size.) 

good hiding-places. 

Bugs, Cicadas, Aphids, and Scale-insects 203 

feeding insects, is found. This difference in food-habit is accompanied 
by more or less obvious structural differences. In the predaceous forms 
the fore legs are usually spined and fitted for seizing and holding the living 
victims, the other legs fitted for swift running, the beak is stout, firm, and 
sharp-pointed, the eyes are often large, protuberant, and flashing bright, 
and there is a general unmistakable air of ferocity about these miniature 
bloodthirsty dragons of the garden shrubbery. 

Five of the terrestrial families of Heteroptera are predaceous, the remain- 
ing eleven being composed of sap-suckers, although in one or two of these 
families a few species seem to have acquired a taste for blood-sucking. 

The largest predaceous family is that of the assassin-bugs, wheel-bugs, 
and soldier-bugs, the Reduviida;. More than fifty genera belonging to 
this family are represented in this country, but so little are the bugs col- 
lected or even noticed by amateurs (or professionals either, for that matter) 
that but few of the species can be said to be at all familiarly known. And 
to use the word "familiarly" in this connection is to indulge in the figure 
of speech known as hyperbole. 

The Reduviids have an unmistakable look of ferocity, small and insig- 
nificant creatures as they are. The eyes are usually large and protuberant, 
looking like a pair of shining black beads set on the small outstretched head. 
The beak, 3-segmented, is strong, sharp-pointed, and large for the small 
head that carries it, and it projects forward in a suggestively eager way. 
While the ground or body color of the bugs is usually black, they are often 
conspicuously marked with blood-red and sometimes with yellow. The 
wingless young are in many species wholly red. A few years ago the news- 
papers were filled with references to a much dreaded "kissing-bug" (one 
of the Reduviids), the name being a satire on the stinging and poisoning 
capabilities of the bug's beak or mouth. The sting, i.e., piercing by the 
beak, of the kissing-bug, and of all other Reduviids, is poisonous because 
of the injection of saliva into the wound, and this poisoning, which makes 
such a wound often very painful and sometimes rather serious to man, must 
be paralyzing and fatal to the more usual insect victims of the assassin-bugs. 
The usual "kissing-bug" of the newspapers is the masked bedbug-hunter, 
Opsicoetus personatus, an insect from ^ to f inch long, blackish brown, 
with prothorax strongly constricted in the middle and longitudinally 
grooved along the middle of the upper surface. The entomologists' 
name for this insect comes from the fact that the young exude a sticky 
substance over the body to which dust, Hnt, etc., adhere so as to cover or 
mask the body, and that the bugs enter houses and prey on bedbugs, cock- 
roaches, and flies. The bite or sting is unusually poisonous and severe. 

Another assassin-bug which forces its acquaintance on us is the "big 
bedbug," or cone-nose, Conorhiniis sanguisugus (Fig. 282), which comes 

204 Bi-igs, Cicadas, Aphids, and Scale-insects 

into houses primarily to drink human blood. It is about an inch long, 
pitchy brown or black, with long narrow head, and with bright red patches 
on the sides of the body and on the base and apex of the fore wings. These 
insects, whose normal outdoors food consists of various insects, often noxious 
ones, as locusts and potato-beetles, are specially common in the South, where 
Comstock says they not infrequently sting children. The banded soldier- 
bug, Milyas cinctus, is a common 
wide-spread friend of the farmer, 
preying on many kinds of noxious 
insects. It is yellow in all stages of 
development with conspicuous fine 
transverse black bands on legs and 
antennae. It roams about over plants 
from early summer to late autumn, 
Fig. 282.-The blood-sucking cone-nose benevolently assimilating the blood 
Conormnus sangmsugiis. (After Howard ■' _ ° 

and Marlatt; natural size.) of its various insect cousins. It glues 

its eggs to the bark" of trees and covers them with a protecting water-proof gum. 
Another fairly well-known member of this family is the wheel-bug, Prionidus 
cristatus, especially common in the South. The full-grown bug is about 
an inch long, black, and has on its thorax a thin convex crest with nine teeth. 
This is the "wheel." The little jug-shaped eggs are laid in six-sided single- 
layered masses of about seventy, which are glued to the bark of trees, or on 
fence-rails, the sides of houses, etc. The young are blood-red, with black 
on the thorax. The wheel-bugs are specially beneficial because they are 
among the few predaceous insects that prey on the well-protected hairy 
caterpillars that infest our shade and orchard trees. 

Closely related to the Reduviids are the curious and readily recognized 
thread-legged bugs, Emesida. The few known species have the body very 
slender and long, and the legs and antennae simply like jointed threads. 
The fore legs, however, are spined and fitted for seizing prey. The common 
species, Emesa longipes (Fig. 283), has the body a little less than i^ inches 
long, each middle and hind leg a little more than ij inches long, and the 
wings when folded not reaching the tip of the abdomen. It is clayey brown 
in color with a reddish tinge above. Howard says that one of the thread- 
legged bugs frequents spiders' webs and robs the spiders of their prey. The 
damsel-bugs (Nabida?) are another small family of predaceous insects which 
usually lurk among flowers and foliage where they capture small insects, 
but which in autumn may often be seen running about on sidewalks and 
elsewhere about houses, probably looking for winter hiding-places. One 
of the commonest and most conspicuous damsel-bugs is the shining jet-black, 
yellow-legged species Corisciis siihcoleoptrahis. The wings and wing-covers 
(in most individuals) are reduced to mere scales, the body is wide and plump 

Bugs, Cicadas, Aphids, and Scale-insects 205 

behind, tapering forward to the narrow prothorax and head. It is about 
^ inch long. The air-bush bug, Phyniata wolfii, a rough, horny-bodied, 
yellowish-green insect with brown or blackish band across the abdomen, 

is about i inch long or less and the body is 
rather like some scaly seed. The abdomen is 
curiously widened behind into two thin, angular, 
scale-like expansions. It conceals itself in 
flower-cups and captures the nectar-sucking 
insect visitors. It is very strong and overcomes 

Fig. 283. 

Fig. 284. 

Fig. 283. — A thread-legged bug, Emesa longipes. (Natural size.) 

Fig. 284. — A damsel-bug, Nabis fusca. (After Bruner; natural size indicated by line.) 

insects, as small butterflies, bees, and wasps, much larger than itself. 

Another small family of blood-sucking bugs is the Acanthiidae, of which 
the most familiar is the wingless degenerate pest, the bedbug, Acanthia 
lectularia (Fig. 285), world-wide in distribution and detestation. To the 
fortunate few who have not at one time or other been forced to a personal 
acquaintance with this bug species it may be told that it is, when full-grown 
and fairly nourished, about \ inch long, reddish brown in color, and broad 
and flat bodied. Small wing-scales or pads can be seen on close examina- 
tion of specimens. The bugs, both immature and adult, can run quickly 
and, because of their "flatness, can conceal themselves in narrow cracks. In 
such crevices in bedsteads, in walls and floors, they hide by day, coming 
out at night to feed. In spring the females lay about two hundred oval 
white eggs in lots of fifty at a time in their haunts in crevices. The eggs 

2o6 Bugs, Cicadas, Aphids, and Scale-insects 

hatch in about a week and the young become full grown in about three months 
moulting five times during growth, but active and capable of "finding" for 
themselves from birth. In the northern states there is but one generation 
a year. The disagreeable bedbuggy odor is produced by a secretion of 
small glands opening, in the adult, on the under side of the body. Another 
species of Acanthia attacks chickens, pigeons, swallows, and bats, and 
Lugger found this species, A hirundinis, or another similar one, attacking 
in daytime the pupils in a school in western Minnesota. The best remedy 
is the free application with a quill-feather of a saturated solution of corrosive 
sublimate (Poison!) in alcohol to all cracks and crevices in infested bed- 
steads, walls, floors, and ceilings. When bedbugs cannot be found hiding 
in bedsteads in daytime and yet mysteriously appear every night, it is often 
because they drop from the ceiling. 

Fig. 2 

Fig. 286. 

Fig. 28s. — The bedbug, Acanthia lectularia; young at left and adult at right. (After 

Riley; natural size indicated by line.) 
Fig. 286. — A predaceous leaf-bug, Lyctocoris fitchii. (After Lugger, natural size 

indicated by line.) 

In this family belong several small inconspicuous insects called flower- 
bugs, which do much good by their persistent preying on noxious insects. 
The best-known species is the insidious flower-bug, Triphleps insidiosus, 
which preys on the chinch-bug. Another species is Lyctocoris fitchii 
(Fig. 286), which preys on the larvae of certain destructive wood-bormg 

The remaining famihes, eleven, of American bugs find their food and 
drink, for the most part, in the juices of living plants. Like the blood- 
sucking bugs, they need for their feeding, and have, a well-developed suck- 
ing-beak. From the tip of the sheath (labium) can be thrust out the foui 

Bugs, Cicadas, Aphids, and Scale-insects 207 

sharp stylets or lancets (maxillae and mandibles) to lacerate the plant-tissues, 
and then the pharyngeal pump sucks up from the wound the flowing sap. 
When too many pumps are drawing away too much sap, the leaves wilt, 
yellow, and die. When too many leaves wilt, the plant starves to death. 
And if the leaves happen to be the corn-leaves, and the pumpers chinch- 
bugs, we have the result estimated for us (by the official U. S. statistician) in 
millions of dollars of loss, as in 1887, when this particular loss in the Missis- 
sippi Valley states was $60,000,000. 

The eleven plant-feeding families of true bugs (Heteroptera) can be 
distinguished by the following key: 

Antennae 4-segmented. 

Fore wings reticulated and of uniform thin substance throughout Tingitid.e. 

Fore wings of various forms or absent, but not reticulated, and not of uniform thin 
substance throughout. 

Beak 3-segmented ; body greatly flattened Aradid.e. 

Beak 4-segmented; body not greatly flattened. 

Membrane (apical area) of fore wings with one or two closed cells at base, but 

otherwise without veins (Fig. 268) Capsid.«. 

Membrane of fore wings with four or five simple or anastomosing longitudinal 
veins arising from the base; or with a larger number of veins arising from 
a cross-vein at the base. 
Ocelli wanting; membrane of fore wings with two large cells at the base, and 

from these arise branching veins (Fig. 268) Pyrrhocorid^. 

Ocelli present. 

Head with a transverse incision in front of the ocelli Berytid^. 

Head without transverse incision. 

Membrane of fore wings with four or five simple veins arising from the 
base of the membrane; the two inner ones sometimes joined to a 

cell near the base (Fig. 268) Lyg.eid^. 

Membrane of fore wings with many usually forked veins springing from 

a transverse basal vein (Fig. 268) Coreid.s:. 

Antennae 5-segmented. 

Scutellum nearly flat, narrowed behind. 

Tibiae unarmed or furnished with very fine short spines Pentatomid^. 

Tibiae armed with strong spines in rows Cydnid^. 

Scutellum very convex and covering nearly the whole of the abdomen. 

Small, black (sometimes with bluish or greenish tinge) CoRiMELiENiDyE. 

Not black Scutellerid^. 

The first of the families in the above table, the Tingitida?, includes the 
curious small lace-bugs (Fig. 287), readily recognized by the dehcate gauze- 
or lace-like appearance of the back, due to the uniform thin and reticulated 
character of the fore wings and of the wing-like expansions of the prothorax. 
About twenty-five species are found in this country, all being plant-feeders, 
living mostly on shrubs and trees. Hawthorn-bushes and oak-, sycamore-, 
and butternut-trees all have particular species of lace-bugs on them. In the 

2o8 Bugs, Cicadas, Aphids, and Scale-insects 

south cotton and beans are also attacked by lace-bugs. The most familiar 
eastern species is the hawthorn lace-bug, Corythuca arcuata, which is com- 
mon on the leaves of hawthorn-bushes. The bugs keep almost exclusively 
on the under side of the leaves. Th'e eggs are laid in small groups on the 
leaves, each egg being imbedded in a little bluntly conical mass of a brown 
sticky substance which hardens soon after egg-laying and looks much like 

a small fungus. The top of the glistening 
white egg can be seen, however, by looking 
down on one of these brown masses. The 
young is broadly oval and flattened in shape, 
brown and spiny, and moults five times in its 
development. The torn, delicate, whitish 
exuviae (cast skins) stick to the leaf. The 
adults hibernate under the fallen leaves 
on the ground beneath the bushes. In 
Cahfornia a similar lace-bug, Corythuca sp., 
(Fig. 287), infests the Christmas berry, 
Heteromeles arhiitijolia, a plant whose clusters 
of bright red berries take the place in Cali- 
fornian Christmas-tide decorations of the 
holly of the East. The eggs (Fig. 287) are 
deposited in the same way as the hawthorn 
lace-bugs', and the life-history is practically 
the same. But because the California winter 
is much less severe and the Christmas berry 
is covered with green leaves all the year, active lace-bugs, young as well 
as adult, can always be found on the bushes. Lace-bugs, small as they 
are, injure any plant on which they gather in numbers, by the continual 
draining away of the sap. Spraying the infested bushes or trees with 
kerosene emulsion (p. 189) will kill the insects. 

The flattest of all the bugs, flatter than the bedbugs even, are the 
curious members of the small family Aradidae. They live in the cracks or 
beneath the bark of decaying trees, and their dull brown color and flat leaf- 
like body make them very difficult to distinguish when at rest in their hiding- 
places. The glistening white eggs are laid under the bark. The flatbugs 
are often mistaken for bedbugs, as they are nocturnal and are often found 
in log cabins. But they probably feed exclusively on plant-sap, being 
especially attracted to mills and recently felled trees, where they suck up the 
sap exuding from the cut or sawed logs. Aradus cinnamomens (Fig. 288) 
is about the same size as a full-grown bedbug and is reddish in tinge, so that 
superficially it does much resemble a bedbug. But all the adult flatbugs 
have wings, while all the bedbugs are wingless. 

Fig. 287. — The lace-bug, Cory- 
thuca sp., of the California 
Christmas berry, Heteromeles 
arbiitijolia; at bottom, eggs on 
small tubercles on leaf; above, 
just-hatched young, intermediate 
stage, and adult. (Eight times 
natural size.) 

Bugs, Cicadas, Aphids, and Scale-insects 209 

The flower-bug family, CapsicUe, contains two hundred and fifty known 
North American species, almost all of which, however, are small and incon- 
spicuous. They mostly five in pastures, meadows, gardens, and along 
roadsides, on the grasses, weeds, and herbaceous flowering plants of these 
places, but some infest woody plants and a few species do much damage 
to garden and orchard shrubs and trees. A few species are predaceous, 
and Howard has seen one species sucking the eggs of the imported elm-leaf 
beetle, a great pest of our elm-trees. The structural characteristic by which 
they can most readily be distinguished from other bugs is the presence of 
one or two closed cells and no longitudinal veins in the membrane (apical 
half) of each fore wing (Fig. 268). When examined closely many of these 

Fig. 288 



Fig. 288.— a flalbug, Aradits cinnamomeus. (After Lugger; enlarged about six times.) 
Fig. 289. — The tarnished plant-bug, Lygus prate.nsis. (Five times natural size.) 
Fig. 29c. — The four-lined leaf-bug, Pmcilocapsus lineatus ; at right, eggs deposited in 

plant-stem. (Figure of insect original, enlarged three and a half times; of eggs, 

after Slingerland, and much enlarged.) 

little bugs will be seen to be elaborately patterned and beautifully colored, 
and their body outhne is trim and graceful. They are active and quick 
to escape from the collecting-net. (The best way to collect them is by sweep- 
ing rankly growing herbage with a short-handled stout net.) Among the 
most abundant and wide-spread Capsids of economic importance is the 
tarnished plant-bug, Lygiis pnitensis (Fig. 289), which attacks many cul- 
tivated plants, as the sugar-beet, strawberry, pear-, plum-, apple-, quince-, 
and other fruit-trees. It is about \ inch long, and ranges from dull dark 
brown to yellowish or greenish brown. A yellowish-white V-shaped mark 
on the scutellum is its most characteristic marking. It hibernates in the 
adult stage, under fallen leaves or in any rubbish, and comes out in the spring 
to pierce and suck sap from tender buds and leaves. The four-lined leaf- 
bug, Poecilocapsus hneatus (Fig. 290), a small bright-yellow bug with head 

2 1 o Bugs, Cicadas, Aphids, and Scale-insects 

and under side of body orange-red, and four black stripes on the back, is 
abundant in the east and north, and is known to attack at least fifty dif- 
ferent kinds of cultivated plants. It is especially familiar as a currant-pest. 
The eggs are deposited in sHts cut lengthwise in plant-stems. The best 
general remedy for these bugs is the jarring of branches of the bushes over a 
dish partly filled with kerosene. Comstock says that the most abundant 
flower-bug in the northeastern states is a small greenish-yellow species with 
two longitudinal black stripes extending from the eyes over the prothorax 
and scutellum. It is long (| inch) and narrow (y^j inch), is found in the 
grass in meadows, and its name is Leptoterna dolobrata. The injury done 
by all Capsids is by the sucking of sap through small punctures and prob- 
ably also, in some cases, the pouring of poisonous saliva into the plant- 
tissues through the punctures. The attacked leaves or buds wilt, turn yellow, 
and finally wither. One of the beneficial Capsids is the glassy-winged bug, 
Hyaliodes vitripennis, a beautiful small yellowish-white insect with almost 
transparent fore wings, with a dash across the apex of these wings, and pro- 
thorax red. It feeds on other insects, and especially on the grape-phyl- 
loxera in its leaf-inhabiting form. Lopidea media is an abundant yellowish- 
red and black Capsid which has learned to like human blood. When it 
cannot have blood it is content with the sap of wild gooseberries. 

The family Pyrrhocoridae is a small family of comparatively large and 
stout bugs, often conspicuous by their colors, of which red and black are 
the most usual. They may be recognized by their 
having the membrane (apical half) of the fore wings 
provided with two large basal cells from which 
several branching veins arise (Fig. 268). They are 
commonly known as "redbugs," and the twenty- 
five species found in our country belong mostly to 
the south and west. The commonest species in 
the north is Largus succindus (Fig. 291), a rusty 
blackish-brown bug about half an inch long, with 
yellowish or pinkish-orange margins on the front 
two-thirds of the back, and a transverse stripe of 
similar color across the base of the prothorax. The 

Fig. 291.— Redbug, Lar- young are steel-blue with a small bright-red dash on 
eus succinctus. (Twice , - ,, , , , , .11, , 

natural size.) DQ.'&e. 01 the abdomen between the backward-pro- 

jecting wing-pads. This species ranges from New 
Jersey to California and south into Mexico. The commonest species of 
the southern states, and one of great economic importance, is the red- 
bug or cotton-stainer, Dysderciis suturellus, which does much damage 
by piercing the stems and bolls of the cotton-plant and sucking the juices, 
but does even more damage by staining the cotton in the opening bolls 

Bugs, Cicadas, Aphids, and Scale-insects 211 

by its reddish-yellow * excretions. Howard says that experiments have 
been made with this insect looking towards its use commercially, and that 
the whole substance of the insect can be converted into a rich orange- 
yellow dye which is readily fixed on woolens or silk by the alum mordant 
liquor. The cotton-stainer is a handsome bug, reddish in color with pale 
brown fore wings striped with pale yellow. The young are bright red with 
black legs. Comstock says that this insect also punctures oranges in 
Florida, so that the fruit begins to decay and drops from the tree. The 
insects can be trapped by laying chips of sugar-cane about the cotton-field or 
orange-grove: the bugs will gather about these chips and may be scalded 
to death. 

One of the largest families of true bugs is the Lygseidae, made notorious 
by a small and obscure representative of it, which, according to the estimate 
of the United States Entomologist, causes our country an annual loss of 
$20,000,000. This insect is the chinch-bug, the 
worst pest of corn, and one of the worst of wheat 
and other small grains. Nearly two hundred species 
of Lygaeids occur in this country, and most of 
them may fairly be called noxious insects. The 
family's structural characteristic most readily noted 
is the presence of but four or five simple longitudinal 
veins in the membrane (apical half) of the fore wings 
(Fig. 268). The antennae rise rather from the under 
than the upper side of the head, and all of the members \' W^^ fJ/ \^ 

of the family have ocelli (simple eyes). While most 
of the Lygaeids are small and inconspicuous, a few fig. 292. — LygcBus turci- 
are comparatively large and bright-colored. The ^"•^- (After Lugger; 
milkweed-bug, Oncopelius fasciatus, about § inch 

long, orange above with most of head and prothorax except the margins 
black, and a broad black band across the middle of the fore wings and 
large black blotch on their tips, is a common showy bug on various 
species of milkweed. An odd-looking, long-necked, common member of 
the family is Myodocha serripes. It is about f inch long, with head long 
and narrow, expanding in front, and rising from a bell-shaped prothorax, 
the rest of the body being elongate and narrow. It is black, with the 
margins, sutures, veins, and some spots on the wing-covers yellow. It is 
common in meadows and thin woods, where it keeps half concealed under 
fallen leaves and twigs. In the south a small species, Pamera longula, \ 
inch long, dark brown with lighter brown on prothorax and fore wings, is 
abundant, feeding mostly on meadow plants. 

Among the many smaller species, the chinch-bug, Blissiis leucopterns 
(Fig. 293), is the best known and most important. It is found nearly all 

212 Bugs, Cicadas, Aphids, and Scale-insects 

over the United States and in Canada, but the great losses occasioned by 
it occur mostly in the corn-growing states of the Mississippi Valley, where 
it has been known as a pest since 1823. I have seen great corn-fields in 
this valley ruined in less than a week, the little black and white bugs mass- 
ing in such numbers on the growing corn that the stalk and bases of the 
leaves were wholly concealed by the covering of bugs. The chinch-bug 
when adult is about ^ inch long, blackish with the fore wings semi-trans- 
parent white and with a conspicuous small trian- 
gular black dot near the middle of the outer margin. 
The very young are red, but become blackish or gray 
as they grow older. The bug is injurious in all 
stages, young, half grown, and adult. The life- 
history, in Kansas, is as follows: The eggs are laid 
in the spring (from middle of March to middle of 
May) by bugs which have hibernated in the adult 
stage. They are laid a few at a time, perhaps five 
hundred in all by each female. The young "red- 
bugs" begin work in the wheat-fields, and usually 

jTjg 2 , 'pj^g chinch- remain in the wheat until harvest (last of June to 

bug, Blissus leucopterus. middle of July), when the destructive host moves into 

(Nine times natural .1 r- u r j • Ti 

gj^g X the fields of young and growmg corn. It requires 

about six weeks for the maturing of the bugs. 

The adults now pair and the cycle of a new generation begins. The 

perfect insects of this generation are those which pass through the winter 

and lay the eggs the following spring for the next year's first brood. It 

is highly probable if not certain that a third brood often appears in Kansas. 

The chinch-bug, though winged, uses its powers of flight but little, and its 

migrations from wheat- to corn-fields in July are usually on foot. The wings 

are used to some degree at pairing-time. 

The remedies for chinch-bug attacks include the gathering together in 

winter of all rubbish, old corn-leaves, dead leaves, etc., in which the old bugs 

hibernate, and burning it, which will destroy many parent bugs, thereby largely 

lessening the spring brood. Disputing the entrance of the bugs into the 

field, when migrating on foot, by plowing furrows around the field and 

pouring coal-tar or crude petroleum into these moats, is often effective. 

There are several natural remedies, namely, the attacks of predaceous insects, 

as aphis-lions, ladybird-beetles, and others, and the attacks of some birds, 

as the common quail. Most effective of all, however, is the rapid spread 

.in a crowded field of a parasitic fungus, Sporotrichiini globtdijerum, which 

kills the bugs by the wholesale. This fungus cannot grow rapidly except 

in moist warm weather, and the bugs thrive especially in dry weather. So 

the rapid spreading and effective killing by this disease depends on favorable 

Bugs, Cicadas, Aphids, and Scale-insects 2 1 3 

meteorological conditions. The "chinch-bug cholera" is well established 
all through the Mississippi Valley, but it can be artificially spread by dis- 
tributing dead and infected bugs in fields where it has not begun to develop. 
This method is followed in several of the corn- and wheat-growing states 
whose entomolgists keep on hand a supply of this fungus — it can be artifi- 
cially cultivated on various nutrient media in the laboratory — to send out 
to farmers on request. The work was begun by Professor F. H. Snow of 
the University of Kansas, and though in the beginning its beneficial results 
were overrated, there is no doubt that much good has come from this wide- 
spread attempt to disseminate artificially the "chinch-bug disease." 

The family Coreidoe, to which the squash-bug, the box-elder bug, and 
certain other more or less familiar insects belong, is another of the larger true 
bug families, being represented in this country by about two hundred species. 
In this family the membrane (apical half) of the fore wings is furnished 
with many veins, most of which arise from a cross-vein near the base (Fig. 
268), and the antennae arise from the upper side of the head. The squash- 
bug, Anasa tristis (Fig. 294), ill-favored and ill-smelling, is a pest of squashes 
and pumpkins all over the country. 
It is brownish black above, with some 
yellow spots along the edges of the 
body, and dirty yellow below. It hiber- 
nates in the adult stage, comes out in 
early spring, and lays its eggs on the 
young sprouts or leaves of squash- and 

pumpkin-vines. The young hatch in f— iiiigKai-^- % mnw n .n 

about two weeks and at first are green, 
but soon turn brown and grayish. 
They suck the sap from the growing Fig. 294. Fig. 295. 

vine, and soon stunt them or even kill Fig. 294.— A squash-bug, Anasa tristis. 
^, ™, ■, ■ . . 4. 4.U (Natural size.) 

them. The remedy is to protect the p^, ap^.-The box-elder bug, Leptocoris 
young plants by means of frames cov- trivittatus. (Twice natural size.) 
ered with netting. After the plants get well started the bugs cannot injure 
them so easily. The box-elder bug, Leptocoris trivittatus (Fig. 295), a con- 
spicuous black insect with three bright-red broad lines on the prothorax 
and the fore wings, with edges and veins of a more dingy red, has become 
familiar with the increased planting of box-elder trees in gardens and streets. 
In the Mississippi Valley and in the plains states these box-elders are much 
used for shade and ornamental trees because of their hardiness, and with this 
increased supply of trees the box-elder bugs have come to be very abundant. 
In late autumn they gather under sidewalks or, often, in stables and houses 
to pass the winter, and have led many housewives to think a new and 
enlarged kind of bedbug had come to town. The bug lives on the sap of 

214 Bugs, Cicadas, Aphids, and Scale-insects 

the trees until winter, and it does not care for much food while hibernating. 
As its mouth is a sucking-beak, it cannot possibly injure hard and dry house- 
hold substances, as some housewives claim. Another Coreid, not uncom- 
mon, is the cherry-bug, Metapodius femoralus, which punctures cherries to 
suck the juice from them. It is dark brown with a rough upper surface, 
and its hind femora are curved thick and knobby, while the hind tibite have a 
blade-like expansion. The leaf-footed plant-bug, Leptoglossus oppositiis, 
is a Coreid destructive to melon-vines, recognizable by the remarkable 
leaf-like expansion of its hind tibiae. A similar leaf-footed species, Lepto- 
glossus phyllopus, occurs in the south, where it attacks oranges and other 
subtropical fruits. 

Allied to the Coreidse is the family Berytidse, or stilt-bugs, of which but a 
few species are known in this country. One of these, Zalysus spinosiis, 
is common all over the country east of the Sierra Nevadas. It is about ^ 
inch long, very slender, and light yellowish brown in color, and is found 
"in the undergrowth of oak woods." Its life-history is not known. 

The remaining four families of true bugs are distinguished by their 
possession of 5-segmented (instead of 4-segmented) antennae (with a few 
exceptions) and by having the body broad, short, and flatly convex, — shield- 
shaped it may then fairly be called, — or very convex or turtle-shaped. Almost 
all of these bugs are exceptionally ill-smelling and have on this account 
got for themselves the inelegant but expressive popular name of stink-bugs. 
As a matter of fact the giving off of offensive odors is characteristic of most 
of the terrestrial true bugs, the squash-bug, chinch-bug, and others being just 
about as malodorous as the so-called stink-bugs. 

Of these four families of shield-bodied bugs, one, the Pentatomidae, is 
represented in this country by numerous species, but the other three con- 
tain but one or two genera each. While most of the Pentatomids, or stink- 
bugs, are plant-feeders, a few are blood-sucking, while some feed indifferently 
on either animal or plant juices. Several of the more common Pentatomids 
are green, as the large green tree-bug, Nezara pennsylvanica, nearly f inch 
long, flattened, with grass-green body margined with a light yellow Une, 
occurring in the fall on grape-vines and other plants; and the bound tree- 
bug, Lioderma ligata, much like Nezara, but with broader body edging of 
pale red and with a pale-red spot on the middle of its back, found 
often abundantly on berries and hazel. Other common stink-bugs are 
brown, as the various species of Euchistes. Still others are conspicuously 
colored with red and black, as the abundant small species Cosmopepla car- 
nijex, about \ inch long, shining black with red and orange spots, most con- 
spicuous of which are a transverse and a longitudinal line in the back of 
the prothorax. The best known and most destructive of these bizarre- 
colored stink-bugs is the harlequin cabbage-bug, or calico-back, Miirgantia 



ugs, Cicadas, Ajmrds^ and Scale-insects 2 1 

histrionica (Fig. 296), black with red or orange or yellow strips and spots, 
which has gradually spread from its native home in Central America to 
all except the northern states of our country. It feeds on cabbages, radishes, 
turnips, and other garden vegetables, and often does great damage in market- 
gardens. In California it has to be fought vigorously in the large market- 


Fig. 296. 

Fig. 297. 


Fig. 296.— The harlequin cabbage-bug, Mur:^anlia histrionica. (Twice natural size.) 
Fig. 297.— The spined tree-bug, Podisus spinosus. (After Lugger; natural length, 

I inch.) 
Fig. 298.— a stink-bug, Pentatoma jtiniperina. (One and one-half times natural size.) 

and seed-gardens of the Santa Clara Valley. The adults hibernate, and in 
the spring each female lays about twelve eggs in two parallel rows on the 
under surface of the young leaves. The young bugs, which are pale green, 
hatch in three days, and in two or three weeks are full grown. There can 
thus be several generations m a season. 

Among the predaceous or blood-sucking stink-bugs the species of the 
genus Podisus are especially common and effective. They destroy many 
injurious insects. Podisus spinosus (Fig. 297), the most familiar species, 
may be recognized by the prominent spine-like processes projecting from 
the posterior lateral angles of the prothorax. The large gray tree-bugs 
of the genus Brachymena with roughened spiny back and grayish body- 
color may be found resting on the bark of trees, with whose color and rough- 
ness they harmonize so thoroughly as to be nearly indistinguishable. They 
feed indifferently on either plant-sap or the blood of other insects. 

Representatives of the three other families of shield-backed or stink- 
bugs will be rarely found by general collectors. The flea-like negro-bug, 
Corimelcena pulicaria (family Corimelffinidffi), is a tiny, very malodorous, 
polished black species often abundant on blackberries and raspberries, 
with which it often goes to market and even farther! The burro wer-bugs 

21 6 Bugs, Cicadas, Aphids, and Scale-insects 

(family Cydnidie) have an oval rounded or elliptical blackish body with the 
front legs more or less flattened and fitted for digging. They are found 
burrowing in sandy places or under sticks or stones. They probably suck 
the sap from plant-roots. 


The members of the suborder Parasita are the disgusting and discom- 
forting degenerate wingless Hemiptera known as lice. They live parasitic- 
ally on the bodies of various mammals, the ones most familiar being the 
three species found on man, all belonging to the genus Pediculus, and the 
several species of the genus Haematopinus found on domestic animals, as 
dogs, horses, cattle, sheep, etc. Both these genera together with a few 
others found on various wild animals, belong to the Pediculidse, the single 
family of the suborder represented in this country. The only other family, 
Polycterridae, contains but two species, both found on bats, one in Jamaica 
and the other in China. 

All the Pediculids are wholly wingless, have the mouth-parts fused to 
form a flexible sucking-tube, and the feet provided with a single strong curved 
claw which specially adapts them for clasping and clinging to hairs. The 

Fig. 299. 

Fig. 300. 

Fig. 299. — The head-louse of man, Pediciilns capitiis. (After Lugger; natural size 

indicated by line.) 
Fig. 300. — The body-louse of man, Pediculus vestimenii. (After Lugger; natural size 

indicated by line.) 

sucking-beak has been described by Uhler as "a fleshy unjointed rostrum 
capable of great extension by being rolled inside out, this action serving 
to bring forward a chaplet of barbs which imbed themselves in the skin to 

Bugs, Cicadas, Aphids, and Scale-insects 217 

gi\-e a firm hold for the penetrating bristles arranged as chitinous strips in 
a long, slender, flexible tube terminated by four very minute lobes which 
probe to the capillary vessels of a sweat-pore." Of the three species of 
Pediculus infesting unclean persons, P. capitus (Fig. 299), the head-louse, 
is longer than wide, whitish with faint dark markings at the sides of the 
thorax and abdomen; P. vestimenti (Fig. 300), the body-louse, is of the 
same shape and general aj)pearance, but when full grown has the 
dorsal surface marked with dark transverse bands; while P. inguinalis 
(Fig. 301), the crab-louse, has the body as wide as long, with strong 
legs spreading out laterally so as to increase the apparent width very 

Fig. 301. 


Fig. 30J 

Fig. 301. — The CTa.b-\ouse.oi ma.n, Phthirius inguinalis. (.'\fter Lugger; much enlarged.) 
Fig. 302. — Egg of crab-louse, Phthirius inguinalis. (After Lugger; much enlarged.) 
Fig. 303. — Sucking dog-louse, HcBmatopinus pilijerus Burm. (After Lugger; natural 
size indicated by line.) 

much. The eggs (Fig. 302), called "nits," of these lice are whitish and are 
glued to the hairs (in the case of P. capitus) or deposited in folds of the 
clothing {P. vestimenti), and the young, when hatched, resemble the parents 
except in size. The whole life is passed on the body of the host. The prime 
remedy for these disgusting pests is cleanliness. Various sulphur and mercu- 
rial ointments will kill the insects. 

The lice of the domestic animals belong to a different genus, Hasma- 
topinus, but are very similar in appearance and structure to the head-lice 
of man. H. pilijerus (Fig. 303), of dogs, is about yV inch long, reddish 
yellow, and with the abdomen thickly covered with fine hairs and minute 
tubercles; H. eiirysternus (Fig. 304), the short-nosed ox-louse, of cattle, 
is from | inch to \ inch long, fully half as wide, with the head bluntly 
rounded in front and nearly as broad as long; H. viiuli, long-nosed ox-louse. 

21 8 Bugs, Cicadas, Aphids, and Scale-insects 

also of cattle, is about ^ inch long and not more than ^ as wide, with long 
slender head, narrow in front; H. urius (Fig. 305), of hogs, is \ inch long, 
being one of the largest of the sucking-lice, with broad abdomen and long 
head, and gray in color, with the lateral margins of head, thorax, and abdo- 

FiG. 304. 

Fig. 305. 

Fig. 304. — Short-nosed cattle-louse, Hamatopinus eiirysternus. (After Lugger; natural 

length 1.5 mm.) 
Fig. 305. — The hog-louse, HcBtnatopinus urius. (After Lugger; natural size indicated 

by line.) 

men black; H. pedalis, the sheep-foot louse, found only on the legs and 
feet of sheep, below the long wool, has a short, wide head and same general 
shape as the short-nosed ox-louse; H. asini, of horses, of about same size 
as the short-nosed ox-louse, but with long and slender head with nearly 
parallel sides; H. spimdosus, of the rat, small, Hght yellow, and with the 
head projecting very little in front of the antennae and the thorax very short ; 
H. acanthopus, of the field-mouse, resembling the rat-louse in color and 
shape, but larger; H. ventricosus, of rabbits and hares, thick-bodied and 
short-legged and with abdomen nearly circular; H. aniennatus, of the 
fox-squirrel, with long slender body and curious curved tooth-like process 
on basal segment; H. sciuropteri, of the flying squirrel, with slender light- 
yellow body, and head as broad as long, and with front margin nearly 
straight; H. siituralis, of the ground-squirrels and chipmunks, with short 
broad golden-yellow body. The eggs of all these forms are glued to the 
hair of the hosts, the young louse escaping by the outer or unattached end 
and immediately beginning an active blood-sucking life. The most effective 

Bugs, Cicadas, Aphids, and Scale-insects 219 

and feasible remedy in the case of thin-haired animals, as swine and horses, 
is the appUcation of a wash of tobacco-water or dilute carbohc acid, or of 
an ointment made of one part sulphur to four parts lard, or kerosene in 
lard, or of a liberal dusting with wood ashes or powdered charcoal; in the 
case of thick-haired animals, as cattle, the best remedy is fumigation by 
enclosing the animal in a sac or tent with the head left free, and burning 
sulphur or tobacco inside the sack. One to two ounces of tobacco and 
exposure of twenty to thirty minutes for each cow have been found effective. 

A BRIEF account of the curious little insects known as thrips may be 
appended here to the chapter on the Hemiptera (Fig. 307). These narrow- 

FiG. 306. 

Fig. 307. 

Fig. 306. — The sheep-louse, Hamatopinus ovis, female and egg. 

size of insect indicated by line; egg much enlarged.) 
Fig. 307. — Thrips, Phorithrips sp. (Much enlarged.) 

(After Lugger; natural 

bodied, fringe-winged, yellowish or reddish brown or blackish little creatures 
can be most readily found in flower-cups, which they frequent for the sake 
of sucking the sap from the pistils and stamens or the delicate sepals 
and petals. Some of them move slowly when disturbed, but others run 
quickly or leap, and nearly all show an odd characteristic bending up o^ 
the tip of the slender abdomen. This movement is usually preparatory 
to flight (in the case of winged individuals), and is believed to be the means 
of separating and combing out the fringes which border both fore and hind 
margins of each wing. There are fine spines on the sides of the abdomen, 
and the movement of the abdomen seems to draw the fringe-hairs through 
these comb-like rows of spines. The thrips vary in size from -^-^ to \ of 
an inch, and may be certainly known by their narrow fringed wings (when 

2 20 Bugs, Cicadas, Aphids, and Scale-insects 

present), which, when the insect is at rest, are laid back along the abdomen 
unfolded, and parallel or slightly overlapping at the tips. Only about forty 
species are yet known in this country, but as practically only one entomol- 
ogist has attempted to make a systematic study of the group and his speci- 
mens were mostly collected in a single locality (Amherst, Massachusetts), 
it is certain that many species are yet to be found and named. This 
entomologist, Hinds, has published in a recent paper (Contrib. to a Mon- 
ograph of the Thysanoptera of N. A., Proc. U. S. Nat. Mus., vol. xxvi, 
1902) practically all that is known of our American species, and I have 
largely drawn on his paper for the present short account. 

Although the thrips used to be classified as a family of the order Hemip- 
tera, they are now, and rightly, assigned to an order of their own, called 
Thysanoptera (fringe-wings). This separation is due tc the peculiar charac- 
ters of their mouth-parts and of the feet, and 
to the interesting character of their develop- 
ment, which is apparently of a sort of tran- 
sitional condition between incomplete and 
complete metamorphosis. The food of the 
thrips is either the sap of living plants or 
moist, decaying vegetable matter, especially 
wood and fungi. The mouth-structure in ac- 
cordance with this food habit is of a sucking 
tvpe, with mandibles and maxilke modified to 
be needle-like to pierce the plant epidermis. 
But the mouth-parts are curiously asym- 
metrical, the right mandible being wholly 
wanting and the upper lip being more ex- 
panded on one side than the other (Fig. 308). 
The peculiarity in the life-history consists in 
a quiescent, non-food-taking stage like the 
pupal stage in insects of complete metamor- 
phosis, but before reaching this stage well- 
developed external wing-pads have appeared, 
just as happens in the case of immature 
insects of incomplete metamorphosis. Finally, 
the peculiar character of the feet is due to the 
presence of a small protrusile or expansile 
membranous sac or bladder at the tip of the 
tarsus, instead of claws or fixed pads, which seero,s to play a not well 
understood function in the holding on by the insect to the leaf or 
flower parts which it may have occasion to visit. The bladder seems 

Fig. 308. — Head and mouth- 
parts, much enlarged, of 
thrips. ant., antenna; lb., 
labrum; tnd., mandible; 7nx., 
maxilla; mx.p., maxillary pal- 
pus; li.p., labial palpus; m.s., 
mouth-stylet, (After Uzel; 
much enlarged.) 

Bugs, Cicadas, Aphids, and Scale-insects 221 

to be expanded by becoming suddenly filled with blood, and contracted 
l)y a receding of the blood. 

The eggs are laid either under bark or on the surface of leaves or, in 
the case of certain species which have a sharp little ovipositor, underneath 
the leaf-epidermis. They hatch in from three to fifteen days, varying with the 
different species observed, and the young grow and feed for from five to 
forty days. Then follows the brief, quiet, non-feeding stage, and the insect 
becomes mature. Probably several generations appear in a year. The 
winter is passed in either larval, pupal, or adult stage, under bark, in dry, 
hollow plant-stems, in lichens or moss, or on the ground under fallen leaves. 
A curious variation in the adults of many species has been noted in reference 
to the wings; adult individuals of a single species may have either fully 
developed wings, very short functionless wings, or even none at all; both 
sexes may be winged, or one winged and the other not; one or both sexes 
may be short-winged or both be wingless. There seems to exist a condi- 
tion somewhat like that in the plant-lice (Aphidida;) , wings being developed 
in accordance with special needs or influences, as scarcity of food, time of 
the year, etc. 

Another peculiarity of the adults is the rarity, and even, apparently, 
the total lack of males in some species. Parthenogenetic development (the 
production of young from unfertilized eggs) is very common throughout 
the order. 

While the food of those thrips most easily found by the beginning student 
is the sap taken from flower parts, most of the sap-drinking species get their 
supply from the leaves of various plants, and when these plants happen to 
be cultivated ones of field or garden, and the thrips are abundant, these 
tiny insects get the ugly name of "pests." Three species in particular are 
recognized by economic entomologists as pests, viz., the onion-thrips 
{Thrips tabaci), the wheat-thrips (Eiilhrips tritici), and the grass-thrips 
(Anaphothrips striatus). The first of these is about ^j inch long, about 
one-fourth as wide as long, and of a uniformly light-yellowish to brownish- 
yellow color. It feeds on many different cultivated plants, as apple, aster, 
blue grass, melons, clover, tobacco, tomato, cauliflower, etc., etc., but its 
chief injuries seem to be to onions and cabbages. It occurs all over Europe, 
England, and the United States, and is probably the most injurious species 
in the order. The wheat-thrips, also but -^^ inch long, brownish yellow 
with orange-tinged thorax, attacks many plants besides wheat, and is very 
fond of puncturing the pistils and stamens of strawberry-flowers, thus often 
preventing fertilization and consequent development of fruit. The life- 
cycle of this species is very short, requiring only twelve days. Eggs depos- 
ited in the tissues of infested plants hatch in three days, the larvae are full- 

222 Bugs, Cicadas, Aphids, and Scale-insects 

grown in five days, and the quiescent pseudo-pupal stage lasts icur days. 
The grass-thrips is the cause of the injury or disease of meadow and pasture 
grasses known as "silver top" or "white top," a common trouble in the 
northeastern states. The male sex seems to be wanting in this species, the 
young all developing parthenogenetically. 



(Order Neuroptera) SCORPION-FLIES 
(Order Mecoptera), AND CADDIS- 
FLIES (Order Trichoptera). 

INN.'EUS, the first great classifier of animals and 
plants, found in the character of the wings a 
simple basis for grouping insects into orders. 
For the wingless insects he established the order 
Aptera ; * the two-winged ones he called Diptera ; 
the moths and butterflies, with scale-covered 
wings, he called Lepidoptera; the beetles with their horny sheath-like fore 
wings he termed Coleoptera; the thin- and membranous-winged ants, bees, 
wasps, and ichneumon-flies he named Hymenoptera; to the roaches, crickets, 
locusts, and katydids, with their parchment-like straight-margined fore wings,' 
he gave the name Orthoptera; the sucking-bugs with their fore wings 
having the basal half thickened and veinless, the apical half membranous 
and veined, he called Hemiptera; and finally he grouped the heterogeneous 
host of dragon-flies, May-flies, ant-lions, lace-winged flies et al., with their 
thin netted- or nerve-veined wings, in the order Neuroptera. 

In the light of our present greatly increased knowledge of the structure 
and development (the two bases of classification) of insects, this primary 
Linnaean arrangement can no longer be accepted as an exposition of the 
true relationships among the larger groups of insects; that is, it is obviously 
not a natural classification. Its greatest faults are that it groups together 
in the Aptera degenerate wingless members of various unrelated groups with 
the true primitively wingless insects, and places together in the Neuroptera 
a host of insects of somewhat similar superficial appearance, but of radically 
dissimilar fundamental structure and development. With increasing knowl- 
edge of the characteristics of the various subgroups in the Linnaean order 
Neuroptera, the too aberrant ones have been gradually one by one removed, 

* Aptera, from a, without, pteron, a wing; Diptera, from dis, double, pteron, a wing; 
Lepidoptera, from lepis, a scale, pteron, a wing; Coleoptera, from koleos, a sheath, pteron, 
a wing; Hymenoptera, from Intmen, a membrane, pteron, awing; Orthoptera, from orthos, 
straight, pteron, a wing; Hemiptera, from hemi, half, pteron, a wing; Neuroptera, from 
neuron, a nerve, pteron, a wing. 

2 24 Nerve-winged Insects; Scorpion-flies; Caddis-flies 

and in most cases given specific ordinal rank. Thus we now consider the 
May-flies to from an order, the stone-flies another, the dragon-flies still 
another, and so on. There are left, grouped together as the order Neu- 
roptera, seven families which possess the common characteristics of netted- 
veined wings (numerous longitudinal and cross veins), mouths with well- 
developed biting or piercing jaws (mandibles), and a development with com- 
plete metamorphosis. Further than this little can be said to characterize 
the order as a whole, and we may proceed at once to a consideration of the 
various distinct families. 


A. Prothorax as long as or longer than the mesothorax and the metathorax combined. 

B. Fore legs greatly enlarged and fitted for grasping Mantispid^. 

BB. Fore legs not enlarged and not fitted for grasping Raphidiid^. 

AA. Prothorax not as long as the mesothorax and the metathorax combined. 

B. Hind wings broad at the base, and with that part nearest the abdomen (the 

anal area) folded like a fan when not in use Sialid^. 

BB. Hind wings narrow at base, and not folded like a fan when closed. 
C. Wings with very few veins, and covered with whitish powder. 


CC. Wings with numerous veins, and not covered with powder. 

D. Antennae gradually enlarged towards the end, or filiform with a 

terminal knob Myrmeleonid^. 

DD. Antennse without terminal enlargement. 

E. Some of the transverse veins between the costa and subcosta 
forked (in all common forms), wings brownish or smoky. 


EE. Transverse veins between the costa and subcosta simple, 

wings greenish Chrysopid^. 

While most of the Neuroptera are terrestrial in both immature and adult 
life, one family, the Sialidse, includes forms whose larvae are aquatic. There 
are only three genera in the family, but all are fairly familiar insects to col- 
lectors and field students. The adults of these genera can be distinguished 
by the following key: 

Fourth segment of the tarsus bilobed; no simple eyes (ocelli) Sialis. 

Fourth segment of the tarsus simple, cylindrical; three simple eyes (ocelli). 

Antennje with segments enlarged at the outer ends; hind corners of the head. rounded. 


Antennae with segments cylindrical; hind corners of the head with a sharp angulation 

or tooth CORYDALIS. 

The larvs can be distinguished by the following key: 
Tip of abdomen bearing a single long, median, laterally fringed tail-like process. .Si.^Lis. 
Tip of abdomen forked, the two fleshy projections each bearing a pair of hooks. 

Lateral filaments (soft, slender, tapering processes projecting from the sides of the abdom- 
inal segments) with no tuft of short hair-like tracheal gills at base. . .Chauliodes. 
Lateral filaments each with a tuft of short, hair-like, tracheal gills at base. .Corydalis. 

Nerve- winged Insects; Scorpion-flies; Caddis-flies 225 

Two species of Sialis occur in this country; they are called alder-flies, 
or orl-flies. The smoky orl-fly, Sialis infitmata, widely distributed over 
this country, is a dusky brownish in- 
sect about ^ inch long, often seen, with 
wings closely folded, sitting on sedge- 
leaves near quiet waters. The larvae 
(Fig. 309), according to Needham, live 
in marshy places tilled with aquatic 
plants, on the borders of streams and 
ponds. When full grown they are 
about an inch long, and keep up an 
undulating motion with the abdomen, 
the long tail being intermittently lashed 
up and down. When full grown the 
larva crawls out of the water and at Fig. 309. — Larva (at right) and pupa (at 

some little distance burrows into the Y!^} °L ^" ""^"^y- . ■^''^^" in/umata. 

. (Alter JNeedham; twice natural size.) 

moist soil for a few inches or even a 

foot or more. Here it forms an oval cell and pupates within it. Two or 

three weeks after the adult fly issues. 

Of Chauliodes, the fish-flies (Fig. 310), eight North American species 

are known. The adults are from i^ to 2^ inches long, and their wings 

expand from 2^ to 4 inches. The wings are grayish or brownish with whitish 

spots or bands, and the antennae are curiously feathered or pectinate. The 

Fig. 310. — The saw-horned fish-fly, Chauliodes serricornis, laying eggs. 
(After a photograph from life by Needham; natural size.) 

larvae live in wet places at the edge of water or in water close to the surface. 
According to Needham they are perhaps oftenest found clinging to the under 
side of floating longs or crawling beneath the loosened bark. They are 
predaceous, feeding upon other aquatic insects. When ready to transform 
they excavate a cell above the level of the water under a stone or log or layer 

226 Nerve-winged Insects ; Scorpion-flies; Caddis-flies 

of moss or in a rotten log, in which they pupate, and from which the adult 

fly issues in about two weeks. 

The genus Corydalis (Fig. 311) is represented by a single species, C. corniita, 

but it is such a conspicuous and wide-spread insect that it is probably the 

best-known species in the whole order 
Neuroptera. The adult fly is most com- 
monly called "hellgrammite," while the 
larvae (Fig. 312), much used by fisher- 
men as bait, are known as dobsons or 
crawlers. But other names are often 
used. Howard lists the following array 
of names, collected by Professor W. W. 
Bailey, which are applied to the larva 
in Rhode Island alone: dobson, crawler, 
arnly, conniption-bug, clipper, water- 
grampus, gogglegoy, bogart, crock, hell- 
devil, flipflap, alligator, Ho Jack, snake- 

FiG. 311. 

Fig. 312. 

Fig. 311. — Dobson-fly, Corydalis corniita, male, with head of female above. (Natural 

Fig. 312. — Larva of dobson-fly, Corydalis corniita. (Natural size.) 

doctor, dragon, and hell-diver. The insect is very common about Ithaca, 
N. Y., and Professor Comstock of Cornell University gives the following 
account of its life-history as observed by him there: " The larvae live under 
stones in the beds of streams. They are most abundant where the water 

Nerve- winged Insects ; Scorpion-liies ; Caddis-flies 227 

flows swiftest. They are carnivorous, feeding upon the nymphs of stone- 
flies, May-flies, and other insects. When about two years and eleven months 

fbnd „^„ 

I ant frf\-~^\ i ,f\^^y\^ 



Fig. 313. — Head of larva, pupa, and adult of dobson-fly, Corydalis cornuta, showing 
development of the mouth-parts of the adult within the mouth-parts of the larva. 
A, head of a larva with its cuticle dissected away on the right-hand side, revealing 
the pupal parts; B, head of male pupa with cuticle dissected away on right-hand 
side, revealing developing imaginal parts; C, head of female pupa with cuticle 
wholly removed, showing imaginal parts; D, head of adult male, md., mandible; 
mx., maxilla; H., labium; lb., labrum; ant., antenna; l.h., larval head-wall; p.h., 
pupal head-wall; ga., galea; li.p., labial palpus; mx.p., maxillary palpus. Any 
of these terms may be prefixed by/, larva; p, pupa; or i, imago. 

old the larva leaves the water, and makes a cell under a stone or some other 
object on or near, the bank of the stream. This occurs during the early 

22 8 Nerve-winged Insects; Scorpion-flies; Caddis-flies 

part of the summer; here the larva changes to a pupa. In about a month 
after the larva leaves the water the adult Insect appears. The eggs are 
then soon laid; these are attached to stones or other objects overhanging 
the water. They are laid in blotch-like masses which are chalky-white 
in color and measure from half an inch to nearly an inch in diameter. A 
single mass contains from two thousand to three thousand eggs. When 
the larvae hatch they at once find their way into the water, where they 
remain until full-grown." 

In the Kansas corn-fields I used to find certain wonderfully beautiful, 
frail, gauzy-winged insects resting or walking slowly about on the great 
smooth green leaves. The eyes of these insects shone like burnished copper 
or shining gold, and this with the fresh clear green (tinged sometimes with 
bluish, sometimes with yellowish) of the lace-like wings and soft body made 
me think them the most beautiful of all the insects I could find. But a 
nearer acquaintanceship was always unpleasant; when "collected" they 
emitted such a disagreeable odor that admiration changed to disgust. These 
lace-winged or golden-eyed flies are common all over the country and com- 
pose a family of Neuroptera 
called ChrysopidcC. All except 
two species of the family belong 
to the single genus Chrysopa, 
which includes more than thirty 
species found in the United 
States. In the Chrysopida; the 
larvae are not aquatic as in the 
family Siahdae, but are active 
and fiercely predaceous little 
creatures called aphis-lions, that 
crawl about over herbage and 
shrubbery in search of living 
aphids (plant-lice) and other 
small soft-bodied insects. The 
aphis-lion (Fig. 314) has a pair 
Fig. 314.— The golden-eyed or lace-winged fly, . , shirn-T^ointed slender 

Chrysopa sp.; adult, eggs, larva, "aphis-lion," ^i long,^ snarp pomtea, sienoer 
and pupal cocoons on the under side of leaf, jaws which are grooved on the 
(Natural size.) -^^^^^ f^^g_ Having found a 

plant-louse it pierces its body with the sharp jaw-points, and holds it up, so 
that the 'blood of its victim runs along these grooves into its thirsty throat. 
The Chrysopa larvae will bravely attack insects larger than themselves, or will 
quite as readily prey on the defenceless eggs of neighbor insects, or indeed of 
their own kind. Indeed, probably because of this egg-sucking habit the female 
lace-winged fly deposits her eggs each on the tip of a tiny slender stem, about 

Nerve- winged Insects; Scorpion-Hies; Caddis-flies 229 

half an inch high, fastened at the base to a leaf or twig (Fig. 314). When 
the first larvae hatch they crawl down the stems and wander around in this 
little forest of egg-trees, but fortunately haven't wit enough to crawl up to 
the still unhatched eggs of their brothers and sisters. When the aphis-lion 
is full-fed and grown, which, in the studied species, occurs in from ten days 
to two weeks, it crawls into some sheltered place, as in a curled leaf or 
crevice in the plant-stem, and spins a small, spherical, glistening, white, 
silken cocoon, within which it pupates. In another ten days or two weeks 
the delicate lace-winged golden-eyed green imago bites its way out, cutting 
out a neat circular piece. 

In the family Hemerobiidas are some insects whose larvae are also called 
aphis-lions; these belong to the typical genus Hemerobius. But in two 
rare genera of the family, Sisyra (Fig. 315) and Climacia, the immature 
stages are aquatic, the small larvae (about i inch long) living as parasites 

Fig. 316. 

Fig. 3156. 

Fig. 315c. 

Fig. 317. 

Fig. 315. — Sisyra umhrata. a, adult; ft, larva; c, pupa. (All about five times natural size.) 
Fig. 316. — Polystwchotes punctatits. (Natural size.) 
Fig. 317. — Hemerobius sp. (Three times natural size.) 

on or in fresh-water sponges (Spongilla). The largest members of the 
family belong to the genus Polystoechotes, of which two species are known. 
The commoner one, P. punctatus (Fig. 316), is about i\ inches long and its 
wings expand 2 to 3 inches. It is nocturnal and is to be collected about 

230 Nerve-winged Insects; Scorpion-flies; Caddis-flies 

lights. Its body is blackish, and the wings are clear but mottled with irreg- 
ular brownish-black spots. When at rest the wings are held steeply roof- 
like over the back. Nothing is known of its life-history. Of the best-known 
genus, Hemerobius (Fig. 317), twenty species have been noted in this country, 
but they are small, dull-colored insects, and are rather rare, or at least 
infrequently seen. Comstock says they occur in forests and especially on 
coniferous trees. The larvae are like the Chrysopa larvae, predaceous and 
well equipped with big strong head and sharp, curved seizing and blood- 
sucking mouth-parts. The larvae (Fig. 318) of some species have the 
curious habit of piling up on their back the empty, shriveled skins of their 
victims, until the aphis-lion is itself almost wholly concealed by this unlovely 
load of rehcts. This is true of all the Hemerobius larvae I have seen in 
California. Stripped of the covering of skins the aphis-lion is seen to have 
a short, broad, flattened body, with numerous long, spiny hairs arising from 
tubercles. These hairs help to hold the mass of insect skins together. 

Still other Neuroptera with fierce, ever-hungry, carnivorous larvae are 
the ant-lions, or Myrmeleonidae. The horrible pit of Kipling's story, into 

Fig. 318. 

Fig. 319. 

Fig 320. 

Fig. 318. — Larva of Hemerobius sp. covered with detritus. (From life; four times natural 

Fig. 319. — Larva of ant-lion, Myrmeleon sp. (Three times natural size.) 
Fig. 320. — Pit of ant-lion and, in lower right-hand corner, pupal sand-cocoon, from 

which adult has issued, of ant-lion, Myrmeleon sp. (About natural size.) 

which Morrowbie Jukes rode one night, is paralleled in fact in that lesser 
world of insect life under our feet. The foraging ant, too intent on bringing 
home a rich spoil for the hungry workers in the crowded nest to watch care- 
fully for dangers in its path, finds itself without warning on the crumbling 

Nerve-winged Insects; Scorpion-tiies ; Caddis-flies 231 

verge of a deep pit (Fig. 320). The loose sand of the pit's edge slips in and 
down, and the frantic struggles of the unlucky forager only accelerate the 
tiny avalanche of loose soil and sand that carries it down the treacherous 
slope. Projecting from the very bottom of the pit is a pair of long, sickle- 
like, sharp-pointed jaws, adapted most effectively for the swift and sure 
grasping and piercing and blood-letting of the trapped victims. The body 
of the ant-lion (Fig. 319) is almost wholly concealed underneath the sand; 
only the vicious head and jaws protrude above the surface in the pit's depths. 
Comstock has seen the ant-lion throw sand up from the bottom, using its 
flat head like a shovel in such a way that the flung sand in falling would 
strike an ant slipping on the slope and tend to knock it down the side. Ant- 
lion pits are to be found all over the country, in warm, dry, sandy places. 
The ant-hons can be brought home alive, and kept in a dish of sand, where 
their habits may be observed. 

The adult ant-lion (Fig. 321) is a rather large, slender-bodied insect 
with four long oar-shaped gauzy wings, thickly cross-veined and usually 
more or less spotted with brownish or black. The eggs are laid in the sand 

Fig. 321. — Adult ant-lion, Myrmeleon. (Natural size.) 

and the freshly-hatched larvae or ant-lions immediately dig little pits. When 
the larvae are full-grown — and just how long this takes is not accurately 
known — each forms a curious protecting hollow ball of sand held together 
by silken threads, lines it inside smoothly with silk, and pupates in this cozy 
and safe nest (Fig. 320). The larva is said to lie for some time, even through 
a whole winter, in this cocoon before pupating. The life-history of no ant- 
lion species is yet thoroughly known. 

The family Myrmeleonidae includes eight genera, which are usually 
grouped into two subfamihes as follows: 

Antennae nearly as long as wings Ascalaphin^. 

Antennae not one-third as long as wings Myrmeleonin^. 

The subfamily Myrmeleoninae includes the true ant-lions with habits 
in general as already described. The live genera in it may be distinguished 
by the following key: 

232 Nerve-winged Insects ; Scorpion-flies; Caddis-flies 

Claws very stout, swollen Acanthaclisis. 

Claws slender at base, not swollen. 

Wings with a black band at tip or eye-like spots Dendroleon. 

Wings not as above. 

Tibia with no spurs (short but conspicuous spines) Maracanda. 

Tibia with spurs. 

Wings with a single row of costal areoles (small cells) Myrmeleon. 

Wings with a double row of costal areoles Brachynemurus. 

The subfamily Ascalaphinse includes but three genera and six species, 
the larvae of which do not dig pits (as far as known), but hide under stones 
sometimes with the body partially covered with sand, or even nearly buried 
in it, and wait for prey to come within reach of their long, sickle-like jaws. 
The adults of this subfamily can be readily recognized by their long antennae, 
knobbed at the tip, like the antennae of butterflies. The habits and life- 
history of Ulula hyalina, an Ascalaphid found in the southern states, have 
recently been studied by McClendon in Texas. The adult fly when at 
rest clings, motionless, to some small branch or stalk, head down with wings 
and antennae closely applied to the branch, and abdomen erected and often 
bent so as to resemble a short brown twig or dried branch (Fig. 322). The 

Fig. 322. Fig. 323. 

Fig. 322. — An Ascalaphid, Ulula hyalina, male. (After McClendon; natural size.) 
Fig. 323. — Larva of Ulula hyalina. (After McClendon; natural size, ^ inch.) 

eggs are arranged in two rows along a stalk and fenced in below by little 
rod-like bodies called repagula, placed in circles around the stalk. The 
eggs hatch in nine or ten days, and the larvae (Fig. 323) crawl down, after a 
day of resting, and hide under stones or in slight depressions. The body 
is covered with sand and the jaws open widely. When a small insect crawls 
within reach the jaws snap together, pinioning the victim on the curved 
points. The jaws are grooved along the inner or lower side and the maxillae 
fit into these grooves so as to form a pair of ducts or channels through which 
the blood is sucked into the mouth. The larva often changes its hiding-place 

Nerve-winged Insects; Scorpion-flies; Caddis-flies 233 

at night. It lives about sixty days, and then seeks a concealed place and 
forms a spherical cocoon of sand and, silk within which it pupates. 

Our three genera of the Ascalaphinae may be determined by the follow- 
ing key: 

Eyes entire Ptynx. 

Eyes grooved. 

Hind margin of wings entire Ului.a. 

Hind margin of wings excised Colobopterus. 

Under the loose hanging strips of bark on the eucalyptus-trees in Cali- 
fornia or on the bark of various Pacific Coast conifers, as pine, spruce, and 
cedar, one may often find certain odd, slender-necked, big-headed, gauzv- 
winged, blackish insects about half an inch long (Fig. 324). A slangy student 
once proposed the name "rubber-neck" for them, and it is a fairly fit one. 
These "rubber-necks," or "snake-flies," belong to the family Raphidiida;, 
of which but two genera are known in the world. The species of the genus 
Raphidia have three simple eyes (ocelli), while those of Inocellia have no 
oceUi. Twenty-four species are found scattered over Asia Minor, Syria, 


Fig. 324. — Raphidia sp., aduh, larva, and pupa. (Two and a half times natural size.) 

eastern Siberia, Europe, and England, while four species of Raphidia and 
three of Inocellia occur in the western half of the United States. The 
snake-flies are predaceous insects, the larvae being notoriously voracious 
insectivores. The larvae live in crevices of bark, or under it, where 
there are breaks in it, as is always the case on old trees of most eucalyptus 

Snake-fly larvae are said to find and eat many larvae of the codlin-moth, 
one of the worst pests of apple-trees. Many of the codlin-moth larvae crawl 
into crevices in the apple-tree bark to spin their cocoon, and there meet 
the hungry snake-fly larvae. 

The pupae (Fig. 324), which are not enclosed in silken cocoons like the 
other terrestrial Neuroptera (ant-Uons, lace-winged flies, Hemerobians), lie 

2 34 Nerve- winged Insects; Scorpion-flies; Caddis-flies 

concealed in sheltered places. They are active, though, when disturbed, and 
look much like the larvae, but are more robust-bodied and bear externally 
the developing wings. The head, with eyes and antennae, is more like that 
of the adult. The complete metamorphosis of these insects seems very 
simple compared with that of such other holometabolous insects as house- 
flies and honey-bees. The adult female (Fig. 324) has a long, slender, 
curved, pointed ovipositor, which probably is used to deposit the eggs in 
deep, narrow, and safe cracks in the bark. But the oviposition has not 
yet been seen, and the full life-history of the Raphidians has yet to be worked 

The extraordinary-looking insect shown in Fig. 325 is one of the few 
members of the Mantispidae, the sixth family of the Neuroptera. Its great 
spiny, grasping fore legs and its long neck make it resemble its namesake, 
the praying-mantis of the order Orthoptera, but its four membranous, net- 
veined wings show its affinities with the Neuroptera. The fore legs are like 
those of the mantis because Mandspa has similar habits of catching hve 
prey with them: it is a case of what is called by biologists "parallehsm of 
structure," by which is meant that certain parts of two animals become 
developed or specialized along similar lines, not because of a near relation- 
ship between them, but because of the 
adoption of similar habits. The wings of 
bats and those of birds show a general 
parallelism of structure, although bats and 
. birds belong to two distinct great groups 
of animals. 

Only two genera, viz., Mantispa and 
Symphasis, of Mantispidae are known, and 

^ ^ , . . ,„ these include but five American species. 

Fig. 325. — Symphasts stgnata. (One . . . \ • r j • 

and one-half times natural size.) Symphasts signata (Fig. 325) IS found m 

California, while of the four species of Man- 
tispa three are found in the East and South, while one ranges clear across the 
continent. But they are insects only infrequently seen, and each captured 
specimen is a prize. The life-history of no one of our species has been studied — 
an opportunity for some amateur to make interesting and needed observations 
— but Brauer has traced the life of the European species, Mantispa styriaca, 
and found it of unusual and extremely interesting character. The following 
account of Brauer's observations is quoted from. Sharp (Cambridge Natural 
History, vol. v): "The eggs are numerous but very small, and are deposited 
in such a manner that each is borne by a long slender stalk, as in the lace- 
wing flies. The larvae are hatched in autumn; they then hibernate and 
go for about seven months before they take any food. In the spring, when 
the spiders of the genus Lycosa have formed their bags of eggs, the minute 

Nerve- winged Insects; Scorpion-flies; Caddis-flies 235 

Mantispa Iarva3 find them out, tear a hole in the bag, and enter among the 
eggs; here they wait until the eggs have attained a fitting stage of develop- 
ment before they commence to feed. Brauer found that they ate the spiders 
when these were quite young, and then changed their skin for the second 
time, the first moult having taken place when they were hatched from the 
egg. At this second moult the larva undergoes a considerable change of 
form; it becomes unfit for locomotion, and the head loses the compara- 
tively large size and high development it previously possessed. The 
Mantispa larva — only one of which flourishes in one egg-bag of a spider — • 
undergoes this change in the midst of a mass of dead young spiders it has 
gathered together in a peculiar manner. It undergoes no further change 
of skin, and is full-fed in a few days; after which it spins a cocoon in the 
interior of the egg-bag of the spider, and changes to a nymph inside its larva- 
skin. Finally the nymph breaks through the barriers — larva-skin, cocoon, 
and egg-bag of the spider — by which it is enclosed, and after creeping about 
for a little appears in its final form as a perfect Mantispa." 

Thus in this insect the larval life consists of two different stages, one 
of which is specially adapted for obtaining access to the creature it is to 
prey on. 

The ConiopterygidcT include a few tiny, obscure insects, the smallest 
members of the order. They have wings with very few cross-veins, and 
both wings and body are covered with a fine whitish powder, hence the name 
"dusty wings" which entomologists apply to them. Only two species are 
known in this country, of neither of which is the life-history known. In 
Europe the larvae of a "dusty wing" species have been found feeding on 
scale-insects. When full-fed these larvae spin a silken cocoon, within which 
they transform. 

The small and little-known order Mecoptera includes certain strange 
little wingless, shining black, leaping insects found on snow, some larger 
net-veined-winged insects with the abdomen of the males ending in a swollen 
curved tip bearing a projecting clasping-organ resembling slightly a scor- 
pion's sting in miniature, and a number of still larger, slender-bodied, narrow- 
winged insects. The only popular name possessed by any of these insects 
is that of scorpion-flies, which has been given the few species with pseudo- 
stings. For these scorpion-flies are not stinging-insects, although the males 
can pinch hard with the caudal clasping-organ. But little is known of the 
life-history of any members of the order, nor is much known of the habits 
of the imagoes. 

There are but five genera in the order, which may be distinguished by 
the following key: 

236 Nerve-winged Insects; Scorpion-iiies; Caddis-flies 

Simple eyes (ocelli) absent. 

Wings well developed; antennae short and thick; body more than J inch long. 


Wings rudimentary; antennae slender; body less than \ inch long Boreus. 

Simple eyes (ocelli) present. 

Abdomen slender, cylindrical; not ending, in males, in swollen tip with clasping-organ. 


Abdomen more robust, and in males conspicuously swollen and curved at tip, and 
bearing pointed clasping-organ. 

Beak elongate, tarsal claws toothed Panorpa. 

Beak short, triangular; tarsal claws simple Panorpodes. 

Boreus is the genus of minute leaping black insects which appear occa- 
sionally in snow. Four species occur in this country, one, B. calijornkus 
on the Pacific coast, two in the northern and northeastern states, and one, 
B. unicolor, found, so far, only in Montana. Of the two eastern species, the 
snow-born Boreus, B. nivoriundus , is shining or brownish black, with the 
rudimentary wings tawny; the other, called the midwinter Boreus, B. 
brnmalis, is deep black-green. Comstock says that both species are found 
on the snow in New York throughout the entire winter, and that they also 
occur in moss or tree-trunks. The females have a curved ovipositor nearly 
as long as the tiny body. Neither their feeding-habit nor life-history is 

The genus Panorpa includes the scorpion-flies, of which fifteen species 
are found in the United States. These insects are from J to f inch long, 
with the wings of about the same length. In all, the body is brownish to 
blackish and the wings are clear but weakly colored with yellowish or 
brownish, and have a few darker spots or blotches, which in one or two 
species cover nearly the whole wing-surface. Part of the head projects 
downwards as a short thick beak, the mouth and jaws 
being at the end. The few observations made on the 
feeding-habits seem to show that the scorpion-flies sub- 
sist mainly on animal matter found dead. They have 

Fig. 326. 

Fig. 326.^A scorpion-fly, Panorpa rufescens. 
Fig. 327. — Larva of scorpion-fly, Panorpa sp. 

Fig. 327. 

(Twice natural size.) 

(After Felt; three times natural size.) 

been seen to attack living injured and helpless insects. Panorpa riijesc 


Nerve-winged Insects; Scorpion-flies; Caddis-flies 237 

(Fig. 326), the commonest species in the eastern states, lays its eggs, accord- 
ing to Felt, in crevices of the ground; the larvas (Fig. 327) hatch in from 
six to seven days and grow rapidly. They burrow in the soil, but not deeply, 
and spend some time wandering about on the surface hunting for food. 
They are full-grown in about one month, probably. The further life-history 
of no American species is yet known, but the larva of a European species, 
when full-fed, burrows deeper in'o the ground, e.xcavates an oval cell in 
a small 'ump of earth and lies in it for several months before pupating. In 
this condition it shrivels to one-half of its previous length, and the body 
becomes curved backwards. If taken out, it moves slowly and cannot 

The species of the genus Bittacus, of which there are nine known in 
our country, are long-legged, slender-bodied, narrow-winged insects (a 
California species is wingless) which do not resemble the scorpion-flies 
much in general appearance, but have a similar 
beak (although longer and slenderer) on the 
head, and have also a similar venation of the 
wings. All the species as far as known are 
predaceous, capturing and eating various kinds 
of insects and probably taking no food except 
that which they catch alive. Bittacus strigosus 
(Fig. 3 8) is the most familiar form in the East. 
I inhabits shady swamps or moist coverts along 
streams, and may be seen restlessly flitting from 
branch to branch, or resting for short times sus- 
pended from a leaf or twig by its long fore legs, 
sometimes by the middle ones also. Its general 
appearance, thus suspended, is not very unlike 
a bit of dried dangling foliage. The position 
appears restful and one might almost think the 
insect asleep. "But it is very far from that," 
says Felt, "as many a small insect could testify 
were it still alive. The small fly that ventures Fig. 
within reach of the long, dangling legs imperils 
its life. In a second those well-armed tarsi seize the unfortunate, the fourth 
and fifth segments of the tarsus shutting together like the jaws of a trap 
with teeth upon their opposing surfaces. The struggle is usually short; 
two, three, or four of those long legs lay hold of the captive and soon 
bring it within reach of the sharp beak. It is only a minute's work 
to pierce a soft part of the body and suck the victim's blood, when 
the lifeless remains are dropped to the ground and the insatiate insect 
is ready for the next." The eggs of this species seem to develop and be 

,328. — Bittacus strigosus. 
(Twice natural size.) 

238 Nerve-winged Insects; Scorpion-flies; Caddis-flies 

dropped a few at a time during the adult life. So far as observed, 
egg-laying consists simply of extruding the eggs and letting them drop at 

The habits of the curious wingless species, Bitiacus apteriis, common 
in California, have been observed by Miss Rose Patterson, a student of 
Stanford University. These long-legged, thin-bodied creatures are not 
readily distinguished among the drying grass-blades where they live, because 
the color of the body is almost exactly like the yellowish tan of the plants. 
Miss Patterson went into the field one windy day when clouds were scudding 
over the sky. At first not a scorpion-fly was to be seen; then, in a brief 
period of sunshine, one was seen swinging itself deliberately along from 
one grass-blade to another. When the wind blew hard it either held firmly 
to the weeds or dropped down to the ground for protection. Finally it took 
up its position near a flower-cluster and clung by all its tarsi. When a bee- 
fly came passing that way it immediately freed two of its legs and held them 
out in an attitude of expectancy. When the fly had passed it remained 
in that position for a minute or so and then relaxed into what seemed a more 
comfortable attitude, holding on by all tarsi. As it became cloudy again, 
the insect dropped down among the weeds and remained near the ground, 
its legs resting on the grass-stems and its abdomen pointing almost directly 
outwards. Miss Patterson disabled a small skipper butterfly and dropped 
it near the Bittacus, but he seemed to pay no attention. A lady-bug did 
not arouse him. A fly passed over and still he did not move. She touched 
him with a pencil-point and he drew back and began to feign sleep. When 
she continued to disturb him he showed an inclination to fight, but did not 
leave his shelter until she forced him to do so by repeated pokes with the 
pencil-point. Then he ran nimbly to the top of a blade of grass and hung 
there: his tarsi went scarcely around the leaves. He remained in that posi- 
tion, motionless, until a bird twittered overhead; then he promptly found 
a sheltered place in a drooping grass-leaf. 

Near him she discovered another scorpion-fly, with a crane-fly in its 
clutches. The crane-fly was still alive and struggled feebly while the scor- 
pion-fly sucked its blood. She disturbed them, but though the scorpion- 
fly stopped its eating, it held its prey as before and moved slowly off with 
it. The body of the crane-fly was almost cut in two by the grasping tarsi of 
its enemy. 

Finding another of the queer creatures swinging on a weed, its four legs 
held out hungrily, she gave it a crane-fly, which it grasped firmly, winding 
the tarsi around its body. The crane-fly struggled, but its captor soon had 
its head buried almost to the eyes in its body. Finally the mangled crane- 
fly gave out. She caught another crane-fly and held it out to the scorpion- 
fly, which thereupon grasped its first victim firmly in one of its hind tarsi 

Nerve-winged Insects; Scorpion-flies; Caddis-flies 239 

and snatched at the second. Then holding both, it began to suck the blood 
of the fresher prey. 

Bringing some scorpion-flies into the laboratory, Miss Patterson placed 
a crane-fly in the jar with a pair of them. The male scorpion-fly seemed 
unusually hungry and soon caught its prey and began to eat. The female 
paid no attention until the male had eaten for some time. Then Miss Pat- 
terson observed the male to bend the posterior portion of its abdomen, and 
between the sixth and seventh and seventh and eighth segments on the 
norsal side of the body rounded organs were quickly protruded and with- 
drawn. Shortly after this the female approached and also began to eat 
the crane-fly. Several times she noted the males attracting the females by 
protruding the "scent-glands." In every case, when the male began to give 
off the scent, the female gradually approached. 

Eggs were laid by the females in the laboratory jars. These eggs were 
pink in color and spherical, although slightly flattened at opposite sides. 
They are simply dropped by the female loosely and singly to the ground. 

In the Rocky Mountains of northern Colorado are some of the most 
attractive "camping-out" places in our land; that is, for "campers" who 
specially like Nature in her larger, more impressive phases. The peaks 
of the Front Range rise to 14,000 feet altitude, and the ice- and water-worn 
canons and great sheer cliffs of the flanks of the Range are only equalled 

Fig. 329. — Phryganea cinerea. (After Needham; enlarged.) 

in this country by the similar ones of the Californian Sierra Nevada. The 
mountain-climber in these wild regions cannot but interest himself in the 
animal and plant life which he finds struggling bravely for foothold in even 
the roughest and most exposed places. To the entomologist the few 
hardy butterfly kinds of the mountain-top, the scarce inhabitants of the 

240 Nerve- winged Insects; Scorpion-flies; Caddis-flies 

heavy spruce forests, and the strange aquatic larvae desperately clinging 
to the smooth boulders and rock bed of the swift mountain streams are 
among the most interesting and prized of all the insect host. So it was 
that my first summer's camping and climbing in the Rockies acquired a 
special interest from the slight acquaintanceship I then made with a group 
of insects which, unfortunately, are so little known and studied in this 
country that the amateur has practically no written help at all to enable 

Fig. 330. — Leptocerus resurgens. (After Needham; enlarged.) 

him to become acquainted with their different kinds. These insects are 
the caddis-flies; not limited in their distribution by any means to the Rocky 
Mountains, but found all over the country where there are streams. But 
it is in mountain streams that the caddis-flies become conspicuous by their 
own abundance and by the scarcity of other kinds of insects. 

In Europe the caddis-flies have been pretty well studied and more than 
500 kinds are known. In this country about 150 kinds have been deter- 
mined, but these are only a fraction of the species which really occur here- 
Popularly the adults are hardly known at all, the knowledge of the group 
being almost restricted to the aquatic larvas, whose cleverly built protecting 
cases or houses made of sand, pebbles, or bits of wood held together with 
silken threads give the insects their common name, i.e., case- or caddis- 
worms. The name of the caddis-fly order is Trichoptera. 

These cases are familiar objects in most clear streams and ponds.. 
Figures 331 and 332 show several kinds. There is great variety in the 
materials used and in the size and shape of the cases, each kind of caddis- 
worm having a particular and constant style of house-building. Grains 
of sand may be fastened together to form tiny, smooth-walled, s^-mmetrical 
cornucopias, or small stones to form larger, rough-walled, irregular cylinders. 
Small bits of twigs or pine-needles may be used; and these chips may be 

Nerve-winged Insects; Scorpion-flies; Caddis-flies 241 

laid longitudinally or transversely and with projecting ends. Small snail- 
shells or bits of leaves and grass may serve for building materials. One kind 
of caddis-worm makes a small, coiled case which so much resembles a snail- 
shell that it has- actually been described as a shell by conchologists. Some 
cases in California streams gleam and sparkle in the water like gold; bits 
of mica and iron pyrites were mixed with other bits of mineral picked up 
from the stream - bed to form 
these brilliant houses. An Eng- 
lish student removed a caddis- 
worm from its case, and pro- 
vided it only with small pieces 
of clear mica, hoping it would 
build a case of transparent walls. 
This it really did, and inside its 
glass house the behavior of the 
caddis - worm at home was ob- 
served. While most of the cases 

are free and are carried about by Ji^^^ \Ol? '-^t^ 

the worm in its ramblings, some Fig. 331. Fig. 332a. Fig. 3326. 

are fastened to the boulders or Fig. 331.— Two cases of caddis-worms. (Natu- 

rock banks or bed of the stream. fig'!^32.-Two cases of caddis-worms with the 
These fixed cases are usually com- larval insects within showing head and thorax 
posed of bits of stone or smooth P'-oJecting. (Natural size.) 
pebbles irregularly tied together with silken threads. In all the cases silk 
spun by the caddis-worm is used to tie or cement together the foreign build- 
ing materials, and often a complete inner silken hning is made. 

Fig. 333. — Halesus indistinctus. (After Needham; enlarged.) 

The larvae within the cases are worm- or caterpillar-like, with head and 
thorax usually brown and horny-walled, while the rest of the body is soft 
and whitish. The head with the mouth-parts, and the thorax with the long 
strong legs, are the only parts of the body that project from the protecting 
case, and hence need to be specially hardened. At the posterior tip of the 

242 Nerve-winged Insects; Scorpion-flies; Caddis-flies 

abdomen is a pair of strong hooks pointing outward. These hooks can 
be fastened into the sides of the case and thus hold the larva safely in its 
house. Numerous thread-like tracheal gills are borne on the abdomen 
and by a constant undulatory or squirming motion of the body a stream of 
fresh water is kept circulating through the case, thus enabling the gills to 
effect a satisfactory respiration. The caddis-worm crawls slowly about 
searching for food, which consists of bits of vegetable matter. Those larvae 
which have a fixed case have to leave it in search of food. Some of them 
make occasional foraging expeditions to considerable distances from home. 
Others have the interesting habit of spinning near by a tiny net (Fig. 335), 

334. — Hydropsyche scalaris. (After Needham; enlarged.) 

fastened and stretched in such a way that its broad shallow mouth is directed 
up-stream, so that the current may bring into it the small aquatic creatures 
which serve these caddis-fishermen as food. The caddis-flies live several 
months, and according to Howard some pass the winter in the larval stage. 
When the caddis-worms are ready to transform they withdraw wholly 
into the case and close the opening with a loose wall of stones or chips and 
silk. This wall keeps out enemies, but always admits the water which is 
necessary for respiration. The pupae in the well-made cases have no other 
special covering, but in the simple rough pebble houses attached to stones 
in the stream they are enclosed in thin but tough cocoons of brown silk 
spun by the larvae. The free cases are also usually attached just before 
pupation to submerged sticks or stones. When ready to issue the pupa 
usually comes out from the submerged case, crawls up on some support 
above water and there moults, the winged imago soon flying away. Some 
kinds, however, emerge in the water. Comstock observed the pupa of one 
of the net-building kinds to swim to the surface of the water (in an aqua- 
rium) by using its long middle legs as oars. The insect was unable to crawl 
up the vertical side of the aquarium, so the observer lifted it from the water 
on a stick. At this time its wings were in the form of pads, but the instant 
the creature was free from the water the wings expanded to their full size 
and flew away several feet. On attempting to catch the specimen Com 

Nerve-winged Insects; Scorpion-flies; Caddis-flies 243 

stock found that it had perfect use of its wings, aUhough they were so recently 
expanded. The time required for the insect to expand its wings and take 
its first flight was scarcely more than one second; certainly less than two. 
As such caddis-flies normally emerge from rapidly flowing streams which 
dash over rocks, it is evident that if much time were required for the wino-s 
to become fit for use, as is the case with most other insects, the wave succeed- 
ing that which swept one from the water would sweep it back again and 
destroy it. 

Fig. 336. 

Fig. 335. — Fishing-net of caddis-worm in stream. (After Comstock.) 
Fig. 336. — Goiiiotatiliiis dispectus. (After Needham; enlarged.) 

The adult caddis-flies are practically unknown to general students. 
They are mostly obscurely colored, rather small, moth-like creatures, that 
limit their flying to short, uncertain excursions along the stream or pond 
shore, and spend long hours of resting in the close foliage of the bank. 
So far as observed the flies take no food, although in all the specimens I 
have examined there are fairly well-developed mouth-parts fitted for lap- 
ping up liquids. They probably do not live long, and certainly do not live 

Fig. 337. — TricBiiodes ignita. (After Needham; enlarged.) 

excitingly. In the Colorado mountains numerous small species occur, 
some w!th beautiful snow-white wings and delicate blue-green bodies (Setodes) ; 
other black-winged, brown-bodied kinds (Mystacides) ; and other light- 
brown winged species (Hydropsyche) in great abundance, but usually the 
adults are comparatively solitary and inconspicuous. They probably fly 

244 Nerve-winged Insects; Scorpion-flies; Caddis-flies 

chiefly at night, as large numbers have been taken in trap lanterns by Betten. 
The eggs are laid, according to this observer, in or directly above the water 
Many clusters of eggs were found under the bark of submerged trees, which 
would lead to the conclusion that in some cases the female insect goes under 
water to deposit the eggs. A spherical cluster found suspended on a sub- 
merged twig under a log floating in deep water contained 450 eggs. 

Some of the caddis-fly larvae can be readily kept in an aquarium. 
Almost any kinds found in ponds will live in aquariums, where their feed- 
ing-habits and transformation may be observed. The caddis-worms that 
build odd cases of small sticks laid crosswise live contentedly in an 
aquarium and are most interesting to watch. The complete life-history 
of no single caddis-fly species has yet been worked out completely, and the 
specific identity of but few of our larvae is known. For three California 
species Geo. Coleman, a student of Stanford University, has obtained adults 
by putting wire-screen cages over the larvae in the streams. In these cages 
the larvae had room enough to hunt food successfully, and they lived, except 
for the circumscribing of their territory, perfectly naturally. Betten has 
similarly reared imagoes from four kinds of larvae in the Adirondack Moun- 

The following keys will enable the collector to classify either his caddis- 
worms (larvae) or caddis-flies (adults) to families: 


Spines on the legs, three simple eyes (ocelli). 

Four spurs on tibiae (second long segment) of middle legs Phryganid^. 

Two or three spurs on middle tibiae Limnephilid^. 

No spines on legs, only hairs or spurs. 

Last two segments of palpi (mouth-feelers) not elongated and flexible. 

Palpi of males 5-segmented; ocelli often present Rhyacophilid^. 

Palpi of males 4-segmented; ocelli absent. 

No spurs on front legs Hydroptilid^. 

Spurs on front legs Sericostomatid.e. 

Last segment of palpi elongate and flexible; palpi hairy. 

Basal segment of antennas long and thick, wings slender, no ocelli. . . .Leptocerid.e. 

Basal segment of antenna shorter, wings broader, last segment of palpi composed 

of numerous subsegments Hydropsychid.e. 

KEY TO FAMILIES {^ARYJE). (After Betten.) 

Larva with head bent downward at an angle with the body; tubercles generally present 
on the first abdominal segment; lateral fringe generally present; gill filaments, 
when present, usually simple. 

Hind legs more than twice as long as the first pair; cylindrical case of sand and small 
stones Leptocerid.e. 

Hind legs not more than twice as long as first pair. 

Nerve-winged Insects; Scorpion-flies; Caddis-liies 245 

Head elliptical, only pronotum (dorsal wall of prothorax) chitinized (horny and 
dark), abdominal constrictions deep; cases of vegetable matter laid longitudi- 
nally and forming a spiral, widening at front end Phryganeid^. 

Head oval to circular, pronotum chitinized, mesonotum often, and metanotum some- 
times chitinized, abdominal constrictions slight. 

Lateral fringe well developed; cases various Limnophilid.e. 

Lateral fringe slightly developed; cylindrical case of sand or small stones. 

arva with head projecting straight forward in line with the rest of body; tubercles 

and lateral fringe wanting; gill-filaments, when present, branched. 
Abdomen much thicker than the thorax; case kidney-shaped, of small stones, or flat 

and parchment-like Hydroptilid^. 

Abdomen little if any thicker than the thorax. 

Third pair of legs a little longer than the first pair; no larval case. .Rhyacophilid^,. 
Third pair of legs about the same length as first pair; no portable larval case. 


THE BEETLES (Order Coleoptera) 

I HE moths and butterflies (Lepidoptera) and the 
beetles (Coleoptera) are the most familiar of the 
insect orders. They are, too, most affected by 
collectors: of all the amateur collectors of insects 
probably nine out of ten collect either Lepi- 
doptera or Coleoptera, or perhaps both. The 
moths and butterflies obviously owe their special 
attractiveness to their beautiful colors and pat- 
terns, and to the interesting metamorphoses 
exhibited in their life-history. A gratifyingly 
increasing number of amateurs and collectors are 
"rearing" or breeding Lepidoptera, and adding much to our scientific knowl- 
edge of them. The beetles owe their place of honor among collectors largely 
to their abundance of species and individuals, the readiness with which 
they can be collected, and the little special attention necessary to their per- 
fect preserv^ation. They are mostly large enough, too, to be handled and 
examined readily, and not so large as to require much cabinet space for 
their keeping. They also make specially fit specimens for exchange. But 
amateurs give almost no attention to the immature stages of beetles. 
Although, like the Lepidoptera, they undergo a complete metamorphosis, the 
larvae are so obscure and usually so concealed underground or in tree-trunks 
or decaying matter or in the water, or, if seen, are so often unattractive and 
even repulsive in appearance — most beetle-larvae are "grubs" — that rearing 
beetles is practically an unknown pastime even with the professed "coleop- 

As a matter of fact, the beetles do not begin to present an interest even 
to professional entomologists at all in proportion to the dominant number 
of species in the order. There is a curious uniformity — with of course the 
startling exceptions which must be mentioned in the same breath with 
almost any generalization about insects — in the general character of the 
structure, development, and habits throughout most of the great order of 
beetles. So that a few life-histories well worked out give us a fair knowledge 

of the principal characteristics of coleopterous development. 





i = Desmocerus palliatus. 
2 = Tragidion armatum. 
3=Chalcophora liberta. 
4= Chrysochus auratus, 
5 = Silpha amftrrivana. 
6=GeotrupJes splendidus 
7 = Chrysochus cobaltinus 
8=Bupr'estis sp. 
9= Calosoma scrutator 
io= Tc-traopes tetraophthahnu; 
ix = Cucujus platipes. 
i2^Meloc sp. 
i3=PeHdnota punctata. 
i4=Parandra brunnea. 
i5=Cyllene robiniae. 
i6=RosaHa funebris. 
i7=Cicindela genetosa. 


!\rayy Jle/lmnn, ,iel. 



It would be reasonable to expect to find the insects of an order so pursued 
bv collectors susceptible of ready classifying and determining. On the 
contrary, no order presents more difficulty to the elementary and even 

labial palpi 

compound eye- 


maxi^la'Ty palpi 








tarsal segme7i{s 


Fig. 338. — Ventral aspect of male great water-scavenger beetle, Hydrophilus sp. 
(Three times natural size.) 

advanced students of systematic entomology. The tables and keys pre- 
pared by the few specialists really competent to determine accurately the 
different species of beetles are as nearly impossible to the amateur and 
elementary student as any "keys" in all the field of classific entomology. 



The characters made use of in separating species, genera, and even families 
are so slight, obscure, and difficult to understand that the tables and keys 
based on them chiefly result in wholly discouraging any beginner who 
attempts to use them. And this is not so much the fault of the systematic 
specialists as of the beetles themselves. When it is recalled that nearly 




^_ ventral nerve 

oviduct'^ ' ^99-ti 
accessory glandi-' Yectuvi 

intestine tubules 

\receptaculum seminalis 
Fig. 33Q. — Dissection of female great water-scavenger beetle, Hydrophihis sp.; 
heart and air-tubes (trachea;) are cut away. (Three times natural size.) 


12,000 species of this order are known in North America north of Mexico; 
that they represent nearly 2000 genera, grouped in 80 families; and that 
much general similarity of structure as well as of habits prevails through- 
out the order, it begins to be apparent whv difficulties in classification are 
inevitable. To find structural differences among these thousands of beetles, 



the specialists have been driven to turn their microscopes on the most obscure 
and insignificant parts of the body, and to take cognizance of the shghtest 
appreciable constant differences. The real way in which an entomologist 
gets his beetles classified is to submit specimens to a specialist for determina- 
tion. Then as his authoritatively determined collection gradually increases, 
the collector begins to get acquainted with certain well-marked species, and 
also with the general appearance or habitus of the members of any one family. 
He becomes in time able to classify his new specimens to families, not bv 
tables or keys, but by general appearance and a certain few characteristic 
structural peculiarities, and to determine some species by comparison with 
the already classified specimens in his collection. The eye thus gradually 
trained becomes more and more discriminating, and the collector may in 
time come to be a recognized "coleopterist" both by virtue of his large col- 
lection and the rare forms it contains and by his wide personal ac- 
quaintanceship with beetle species. In the necessarily limited account of 
the Coleoptera given in the following pages I purpose to give keys only to 
tribes and families, and, in order to make even these simple enough to be 
useful, to leave most of the small, rare, and obscure families wholly out of 

The tables thus freed of over half the families of the order still include 
five-sixths of all the North American beetle kinds, and will be found to include 
nine out of every ten beetle species collected. That is, the great proportion, 
ninety per cent, probably, of species at all common enough to be collected 
belong to less than half of the recognized families. These more familiar 
families can also be grouped into a few tribes, each having some simple 
common structural characteristic, thus still further aiding in the work of the 
classifier. The collector will thus first classify his specimen to a tribe by 
means of the table on page 251, and then turning to a discussion of that 
particular tribe find a key to its families.* In the discussion of each of 
these will be found accounts of the life of certain of the more abundant, wide- 
spread, and interesting species of the family. 

The characteristics of the order as a whole are obvious and familiar: 
most beetles are readily known for beetles, and but few insects of other orders 
get mistaken for them. The "black beetle" of the house is a cockroach, 
and several of the hard-bodied, blackish sucking-bugs are sometimes mis- 
takenly called beetles, as are also the earwigs. But the horny fore wings, 
elytra, serving as a sheath for the large membranous hind wings, the true 

* If the collector wishes a further determination of his specimens, he must do as prac- 
tically all other amateur and most professional entomologists do; that is, send his 
material to a specialist, who has, by the way, the right recognized by custom of keeping 
any of these specimens sent him, to add to his own cabinets, ft is well, therefore, to 
send an extra specimen to return in the case of any species likely to interest him. 



organs of flight; the firm, thick, usually dark, chitinized cuticle or outer 
body-wall; the strong-jawed biting mouth, and the compact body, usually 
short and robust, are structural characteristics obvious and usually dis- 

FiG. 340. — The different forms of antennae of beetles, i, serrate; 2, pectinate; 3, cap- 
itate (and also elbowed); 4-7, clavate; 8-9, lamellate; 10, serrate; 11, irregular 
(Gyrinus); 12, 2-segmented antennse of Adranes ccbcus. (After LeConte.) 

tinctive. Especially used in classification are the differences in number 
of tarsal segments of the feet, and differences in the character of the antennae. 
To learn the range of these differences in the antennae, and the names applied 
to the various kinds a careful inspection of Fig. 340 will do more than a 

Fig. 341. — Different forms of legs and tarsi of beetles. (After LeConte and Comstock.) 

page of description. Similarly Fig. 341 illustrates the range of the charac- 
ters drawn from the tarsi. 

The development of beetles is "with complete metamorphosis "; that is, 
from the eggs laid underground, or on leaves or twigs, in branches or trunks 
of live trees, in fallen logs, on or in decaying matter, in fresh water, etc., 



hatch larvae usually called grubs, with three pairs of legs (sometimes want- 
ing), with biting mouth-parts, simple eyes, and inconspicuous antennae. 
These larva2 are predaceous, as the water-tigers (larvae of water-beetles), 
plant-feeders, as the larva? of the long-horns, or carrion-feeders, as those of the 
burying-beetles, and so on. They grow, moult several times, and finally change 
into a pupa either on or in the food, or very often in a rough cell under- 
ground. From the pupa issues the fully developed winged beetle, which 
usually has the same feeding-habits as the larva. The special food-habits 
and characteristics of development are given for numerous common species 
in the accounts (postea) of the various more important families of the order. 

The enonomic status of the order Coleoptera is an important one. So 
many of the beetles are plant-feeders, and are such voracious eaters in both 
larval and adult stages, that the order must be held to be one of the most 
destructive in the insect class. Such notorious pests as the Colorado potato- 
beetle, the two apple-tree borers, round-headed and flat-headed, the "buffalo- 
moth" or carpet-beetle, the wireworms (larvae of click-beetles), the white 
grubs (larvs of June beetles), rose-chafers, flea-beetles, bark-borers and 
fruit- and grain-weevils, are assuredly enough to give the order a bad name. 
But there are good beetles as well as bad ones. The little ladybirds eat 
unnumbered hosts of plant-lice and scale-insects; the carrion-beetles are 
active scavengers, and the members of the predaceous families, like the 
Carabids and tiger-beetles, undoubtedly kill many noxious insects by their 
general insect-feeding habits. 

The great order Coleoptera is divided into two primary groups, some- 
times called suborders, namely, Coleoptera genuina, the typical or true 
beetles, including those species in which the mouth-parts are all present and 
the front of the head is not elongated into a beak or rostrum, and the 
Rhynchophora, snout-beetles (p. 294), which have the front part of the 
head more or less extended and projecting as a beak or rostrum, and the 
mouth-parts with the labrum (upper Up) so reduced as to be indistinguish- 
able and the palpi reduced to mere stiff jointless small processes. To 
this latter suborder belong those beetles familiarly known as weevils, bill- 
bugs, bark-beetles, and snout-beetles. 


With five tarsal segments in all the feet (with rare exceptions). Section Pentamera. (p. 252). 
With the antennae slender, thread-like, with distinct, cylindrical segments. 

(Carnivorous beetles.) Tribe Adephaga (p. 252). 
With the antennas thickened gradually or abruptly toward the tip. 

(Club-horned beetles.) Tribe Clavicornia (p. 258). 
With the antennae serrate or toothed. 

(Saw-horned beetles.) Tribe Serricornia (p. 265). 

With the antennae composed of a stem-like basal part, and a number of flat blade-like 

segments at the tip. (Blade-horned beetles.) Tribe Lamellicornia (p. 272). 




With four tarsal segments in each of the feet Section Tetramera (p. 277). 

Mostly with slender cylindrical antennte, sometimes very long and thread-like, 
sometimes shorter and thickened toward the tip; the fourth and fifth seg- 
ments of the tarsus closely fused, the fourth segment being very small and 
sometimes difficult to distinguish. 

(Plant-eating beetles.) Tribe Phytophaga (p. 277). 

With three tarsal segments in each of the feet Section Trimera (p. 286). 

With the front and middle legs with 5-seginented tarsi, and the hind legs with 4-seg- 
mented tarsi Section Heteromera (p. 288). 


In the tribe of Adephaga, or carnivorous beetles, are four principal 
families, which may be distinguished by the following key: 

Antennae inserted on front of the head above the base of the mandibles. 

(Tiger-beetles.) Cicindelid^. 

Antennce inserted on side of the head between the base of the jaws and the eyes. 

(Predaceous ground-beetles.) Carabid^e. 

With two eves (Predaceous diving-beetles.) Dytiscid^. 

With four eyes, two above and two below (Whirligig-beetles.) Gyrinid^. 

The attractive tiger-beetles (Cicindelidae) are great favorites with col- 
lectors, and deservedly. Their vivid, sharply marked metallic colors, trim 
clean body, and constant alertness and activity, together with their fond- 
ness for warm, bright hunting-grounds and their clever and "gamy" 
elusiveriess of the collecting-net, combine to give these 
fierce, swift little creatures a high place in the regard of 
the beetle-catching sportsman. There are but four genera 
in the family, but the genus Cicindela contains about 
sixty species, distributed over the whole country. In 
California we are not provided with quite our share of tiger- 
beetles, but then there are not so many Cicindelid-hunters 
as in the East. Look for tiger-beetles on sunny days in 
hot dusty roads or open sandy spots. In cold and cloudy 
weather, and at night, they lie hidden under stones or 
chips or in burrows, although a few species are nocturnal 
in habit. When out and running or flying about they are 
hunting; their big eyes and long sharp mandibles and the 
whole seeming of the body some way betray their predatory 
habits even before one sees the swift pounce on some 
dull-witted, slow-footed insect, and the eager blood- 
drinking immediately thereafter. 
The egg-laying habit of the tiger-beetles is not yet known, but the larvae 
and their habits are familiar. They are ugly, malformed, strong-jawed 

Fig. 342. — Larva 
of a tiger- 
beetle, Cicindela 
hybrida. (After 
Schiodte ; three 
times natural 


TIGER BEETLES. (After Lcng and Bcutenmuller.) 

Ig. I. Tctracha Carolina. 

" 2. Cicindela unipunctata 

" 3- 

' celeripes. 

" 4 

' dorsal is. 

" 5 • ' 

' scutellaris var. rugifrons. 

" 6 

' longilabris 


' " var. pcrviridis 

" 8 

' scutellaris var. Lecontei 

" 9 

' sexguttata. 

" lO ' 

' " var. patruela. 

" II ' 

' purpurea. 

" 12 ' 

" var limbalis. 

" 13- 

' formosa var. generosa. 

" 14 

' aneocisconensis. 

" IS 

' vulgaris. 

" i6 

' repanda. 

" 17. 

" 12. guttata. 

" 18 

' hirticollis. 

" 19. 

' punctulata. 

"20 ' 

' marginata. 

" 21 ' 

' puritana. 

" 22 ' 

' lepida. 

" 23 

' rufiventris. 

" 24 

' Hentzii. 

" 25. 

' tortuosa. 

" 26 

' abdominalis. 

" 27 

' marginipennis. 


I |i I I I 


( i 









I' I 



grubs (Fig. 342) which lie in the mouth of a vertical burrow several inches 
deep, with the dirt-colored head bent at right angles to the rest of the body 
and making a neat plug for the top of the hole. When an unwary insect 
comes in reach of this plug the waiting jaws make a quick grasp, and the 
doomed prey is dragged down into the darkness. On the fifth segment 
of the abdomen of the larva there is a hump, and on it are two small but 
strong hooks curved forward. "This is an arrangement by which the little 
rascal can hold back and keep from being jerked out of its hole when it gets 
some large insect by the leg, and by which it can drag its struggling prey 
down into its lair, where it may eat it at leisure. It is interesting to thrust 
a straw down into one of these burrows, and then dig it out with a trowel. 
The chances are that you will find the indignant inhabitant at the remote 
end of the burrow chewing savagely at the end of the intruding straw." 

Plate III shows the appearance of the body and the character of the mark- 
ings of the tiger-beetles, while the vivid color-effects are illustrated in Plate 11. 
In the East occurs, besides Cicindela, the genus Tetracha (PL III, Fig. i) 
with two species; on the plains of the middle West the largest member of 
the family, Amblychila cylindriformis, which hunts its prey at twilight, and 
on the Pacific coast the genus Omus with ten species, all nocturnal. 

The family Carabidae, the predaceous ground-beetles, is a large one, 

including in North America about 1200 species, representing over a hundred 

genera. They are mostly dark-colored and are nocturnal in habit, hiding 

by day under stones, chips, logs, etc., so not many of them are familiar or 

even often seen. A few, however, are large and brilliantly colored, and 

get discovered by most collectors. Like the tiger-beetles 

they are active and predatory, with long strong mandibles 

and slender running legs. They differ from the tiger-beetles 

in their dislike of daylight, and in having the head in 

most species narrower than the thorax. The larvae (Fig. 

343) are "mostly long flattened grubs with a body of almost 

equal breadth throughout. It is usually protected on top 

by horny plates and ends in a pair of conical and bristly 

appendages." Most of the larvtC burrow just beneath the 

surface of the earth, feeding on various insects which enter 

the ground to pupate or for other reasons. They destroy 

large numbers of the destructive leaf- feeding beetles, whose ; 343-— Larva 

° . ° ' 01 Lalosoma sp. 

soft-bodied larvae leave the plants and burrow into the (After Lugger; 

ground when ready to pupate. When full-grown the Carabid enlarged.) 
larvae form small rough cells in the soil within which they change to pupae. 
When the adult beetles emerge they push their way up to the surface. 

Plate IV illustrates several species of this family and shows the charac- 
teristic flattened, usually rather broad although trim and compact, shape 



of the body. In most of the species the elytra are marked with fine longi- 
tudinal lines or rows of punctures, and in several species the hind wings are 
wanting, so that flight is impossible. There is something characteristic 
and almost unmistakable about the general make-up and appearance of 
these beetles. Their flatness, and smoothness, their shining black, greenish, 
or brownish coloration, and their small head with prominent, projecting, 
slender antennae, pointed mandibles, conspicuous clubbed palpi, and bright 
eyes, together with their equally characteristic haunting of hidden places 
on the ground, their swift alert running, and readiness to bite when caught, 
distinguish them, almost at a glance, from all other beetles. One of the 
largest, most conspicuous and well-known Carabids is the searcher, or cater- 
pillar-hunter, Calosoma scrutator (PI. II, Fig. 9), an inch and a half long, 
with vivid violet-green elytra margined with reddish. It is commonly found 
at twilight and after dark on trees, and is often seen by collectors when 
"sugaring" for moths. It is said to make special war on the hairy tent- 
caterpillars, and thus do much good. Two other species of this genus, 
C. jrigidum (Fig. 344) and C. cahduni (Fig. 345), the latter 
called the fiery hunter from its characteristic rows of reddish 
or copper-colored punctures on the black elytra, are keen 

Fig. 346. 

Fig. 344. Fig. 345. 

Fig. 344. — Calosoma frigidum. (After Lugger; natural size.) 
•fiG. 345. — Calosoma calidum. (After Lugger; natural size.) 
Fig. 346.— Larva of Pterostichus striola. (After Schiodte; two and one-half times natu- 
ral size.) 

hunters of cutworms, canker-worms, etc. At the other extreme of size 
in the family are the tiny Bembediums and Tachys, some species of 
which are but yV inch long. The curious bombardiers, or bombarding 
beetles (Brachina), when disturbed, spurt out with popgun sound and puff 
of "smoke" an ill-smelling, reddish, acid fluid from the tip of the 
body. Comstock says that "these beetles have quite a store of ammuni- 
tion, for we have often had one pop at us four or five times in succession 





Fig. I 

Panagsus f 


" 2. 



" 3- 



" 4. 

( ( 


" 5- 

( ( 


" 6. 



" 7- 



" 8. 



" 9- 



" lO. 



'* II. 






while we were taking it prisoner." These beetles have a narrow reddish- 
yellow head and prothorax, and blackish-blue elytra. Of similar appear- 
ance is Lebia grandis, the enemy of the Colorado potato-beetle, feeding 
on its egg and larvoe. Most abundant of the Carabids are the numerous 
dull-black medium-sized species of Pterostichus (PI. IV), in which the pro- 
thorax has a narrow, flat, projecting margin. Over one hundred species 
of this genus have been found in this country. Harpalus is another large 
genus with some very common species; H. pennsylvanicus is often found 
in orchards eating the larvae of the codlin-moth and plum-curculio, ravag- 
ing fruit-pests. A few Carabids are not such good friends, Lugger record- 

FiG. 347. — Predaceous diving-beetles (and back-swiramers, order Hemiptera) in water. 
(From life; slightly less than natural size.) 

ing the fact that Agonoderus pallipes, a species abundant in Minnesota, 
sometimes feeds on sprouting seeds of corn. 

Predaceous beetles of very different habitat are the Dyticidae, the carniv- 
orous water- or diving-beetles. Three hundred species occur in this country, 
and some members of the family are to be found wherever there are streams 
and ponds. They vary in size from the large Cybister and Dyticus, an 
inch and a half long, to small species of Hydroporus and other genera less 
than a fifth of an inch long, but all are readily distinguishable from their 
aquatic companions, the whirligigs (family Gyrinidae) (p. 257), by having 
but one pair of eyes, and from the water-scavenger beetles (family Hydro- 




philidae) (p. 258) by having slender thread-like antennae instead of clubbed 
ones. All are oval and flatly convex in shape, w^ith hard smooth body-wall, 
usually brownish or black, and when at rest hang head downward from 
the surface of the water, the characteristic breathing attitude. The females 
sometimes have the elytra furrowed with shallow longitudinal grooves, and the 
males of most species have a curious clinging-organ on the expanded first three 
or four tarsal segments of the front feet (Fig. 349). This organ is com- 
posed of a hundred or more small capsules on short stems and two or three 
very much larger pads. It is used for holding the females in mating, and 
adheres to their smooth body-wall by the secretion of a gummy fluid insol- 
uble in water. The pads and capsules may also act to some extent as 
"suckers" by atmospheric pressure. The hind legs are long, strong, 
and flattened to form oars or swimming-organs. This beetle regularly and 
perfectly "feathers its oars" by a dexterous twist while swimming. To 
breathe, the beetle comes to the surface — its body being less dense than 
water, it floats up without effort — and projects the tip of its abdomen through 
the surface film. It now lifts the tips of the elytra slightly; air pours in 
and is held there by the fine hairs on the back, where are also the spiracles, 
or breathing-openings. Thus when the beetle goes down 
again it carries with it a supply of air by means of which 
respiration can go on for some time under water. The 
diving beetles can be readily kept in aquaria, as can also 
their larvag (described in the next paragraph), and the 
interesting active Hfe with the characteristic swimming, 

diving, breathing, captur- 
ing of prey, and feeding 
all easily observed. 

The life-history of 
no American species has 
been completely worked 
out, but the eggs of some 
species are dropped ir- 
jv^ regularly on the water, 
while those of others are 
laid in slits cut by the 
sharp ovipositor of the 
female in the stems of 
aquatic plants. The long, 
slender, semi-transparent, 
predaceous larvae (Fig. 348) are known as water-tigers. They have six slender 
legs and the head is large and flattened. It bears long, slender, curved, 
sharp-pointed, hollow mandibles, each with a small opening at the tip and 

Fig. 349. 

Fig. 348. — Water-tiger, the larva of the predaceous water- 
beetle, Dyticus sp. (Natural size.) 

Fig. 34Q. — The predaceous water-beetle, Dyticus sp., pupa 
and adult. (Natural size.) 



another near the base. When a Hve insect or other aquatic creature is caught 
by the active larva its body is pierced by the mandibles and the blood sucked 
through them into the mouth, the opening at the base just fitting, when the 
mandibles are closed, into the corners of the small silt-like mouth. Both 
larvae and adults are fierce and voracious, and the larger species attack and 
kill small fish. In the middle states these beetles actually do much damage 
in cfarp-ponds. The larva breathes through a pair of spiracles at the slender 
tip of its body, which is thrust up to the air when it comes to the surface 
of the water. When ready to pupate it leaves the water — breathing now 
also through six pairs of lateral spiracles — and makes a rough cell in the 
ground of the pond or stream bank. "The pupa state lasts about three 
weeks in summer; but the larvae that transform in autumn remain in the 
pupa state all winter." 

The larger of our common species belong to Cybister, Dyticus, and 
allied genera. In Cybister the little cups on the under side of the tarsal 
disks of the male are similar, and arranged in four rows. In Dyticus and 
its allies the cups of the tarsal disks vary in 
size. Fig. 349 represents a common species of 

" The most common of the diving-beetles 
that are of medium size belong to the genus 
Acilius. In this genus the elytra are densely 
punctured with very fine punctures, and the 
females usually have four furrows in each wing- 

An interesting account of the habits 
and special structures of the common large 
European diving-beetle, Dyticus marginalis, 
is given in Miall's Natural History of Aquatic 
Insects, pp. 39-61. 

Smaller than the predaceous diving-beetles, 
and readily recognized by their curious spin- 
ning or circling, in companies, on the surface 
of ponds or still pools in streams, are the 
whirligig-beetles (Gyrinidffi), common all over 
the country. About forty species of these 
beetles, varying in size from one-sixth to three-fourths inch in length, have 
been found in North America, three-fourths of them belonging to the genus 
Gyrinus. They are all of similar shape and steely blue-black in color, 
and have the compound eye, on each side, wholly divided into an upper 
and a lower part by the sharp lateral margin of the head. Like the 
Dyticids, the whirligig-beetles breathe at the surface and carry air down with 

Fig. 350. Fig. 351. 

Fig. 350. — Whirligig - beetle, 
Dineutes emarginata. (Twice 
natural size."! 

Fig. 351. — Larva of whirligig- 
beetle, Gyrinus marinus. 
(After Schiodte; enlarged.) 




them when diving or swimming below the surface, by having a bubble 
attached to the posterior tip of the body. The hindmost legs are broad 
and paddle-shaped, and fringed with long stiff hairs. The whirhgig- 
beetles can fly, but usually have to climb up on some weed or stick pro- 
jecting from the water in order to make a start. They can make a curious 
squeaking noise, probably a call to other whirligigs, by rubbing the under 
side of the wing-covers against the end of the body. When handled, most of 
these beetles emit an ill-smeUing whitish hquid. 

In the winter the whirligigs lie torpid in mud among the roots of water- 
plants, coming out by twos and threes in the spring. The eggs are laid 
usually on the leaves of some water-plant, and the curious slender larva 
(Fig. 351) is provided with long tapering lateral gills fringed with fine hairs. 
There is a pair of gills on each abdominal segment. It feeds on water- 
insects and other small aquatic animals, and probably also on the "tender 
parts of submerged plants." The pupae of but few species are known. 
That of a common English species lies in a grayish silken cocoon spun on 
some water-plant above the water's surface. 


The clavicorn beetles, or those with clubbed antennae, show much variety 
in the character of the terminal thickening of the antennae (Fig. 340, 4-7), 
which is the characteristic structural feature of the members of the group, 
and from which the tribal name is derived. The tribe includes, too, beetles 
of widely different habits, some aquatic, others terrestrial, some predaceous, 
others plant-feeding, others living on dry stored grains, woolens, and still 
others feeding on carrion. They have indeed little in common and the 
grouping is largely a matter of convenience in classifying. The more im- 
portant families of this tribe can be separated by the following key: 

Aquatic; legs fitted for swimming (Water-scavenger beetles.) Hydrophilid^. 

Terrestrial; legs not fitted for swimming. 

Antennae moniliform, i.e., with segments bead-like; elytra usually covering only basal 

half of abdomen (Rove-beetles.) STAPHYLiNiDiE. 

Antennae moniliform or sub-moniliform; elytra covering most of the abdomen: brown 

or reddish species (Grain-beetles, etc.) Cucutid^. 

Antennas capitate, i.e., ending in a little ball, or clavate. 

Large insects, the smaller not much less than half an inch long (except Catops); 

body usually flattened (Carrion- or burying-beetles.) Silphid^. 

Small insects, mostly less than one-half inch long; body thick and convex above. 

(Larder-beetles, etc.) Dermestid^. 

In the same ponds and pools with the predaceous diving-beetles and 
whirligigs may be found other water-beetles, black, shining, and often of 
large size, which are readily distinguished by their short concealed clavate 



antennae (the long slender palpi may be at first glance mistakenly taken 
for antennae) as members of the family Hydrophilidce, the water-scavenger 
beetles. As the popular name indicates, these beetles feed, for the most 
part, on decaying material, animal or plant, found in the water, although 
they feed also on Uving water-plants, as Nitella; and living insects are cer- 
tainly taken by some species. They can be distinguished from the Dyticida) 
when swimming by their use of the oar-legs alternately, and when at the 
surface getting air by hanging there head upward. The air spreads in a 
thin silvery layer over the ventral side of the body, held there by fine pubes- 

The eggs are deposited in a ball-like silken cocoon with a curious handle- 
like tapering curved stem or spike (Fig. 353). The cocoon floats freely 
on the water, or is attached to some floating leaf or grass-blade or stem. 
From fifty to a hundred eggs are enclosed in each sac. The larvae (Fig. 
354) are elongate, but thicker and less graceful than 
the water-tigers (larvae of the Dyticidae), and, unlike 
the adults, feed chiefly on living insects, snails, tad- 

Fig. 352. Fig. 353. Fig. 354. 

Fig. 352.^Great water-scavenger beetle, Hydrophilus triangularis. (Natural size.) 
Fig. 353. — Egg-case of great water-scavenger beetle, Hydrophilus sp. (Twice natural 

Fig. 354. — Larva of great water-scavenger beetle, Hydrophilus caraboides. (After 

Schiodte; natural size.) 

poles, etc. They breathe through spiracles at the tip of the body, coming 
occasionally to the surface to get air. In shallow water they simply lie 
with the tip of the tail projected up to the surface. When ready to pupate 
the larvae leave the water, and, burrowing a few inches into the ground, form 
a rough cell in which they transform. The adult beetles fly readily, and 
sometimes, with Dyticids, are to be found at night around electric lights. 
When winter comes they burrow into the bottom or bank of the pond or 
stream and lie torpid until spring. 



About one hundred and fifty species of HydrophilidEe are known in this 
country. The largest species belong to the genus Hydrophilus, are shining 
bluish or greenish black, and measure nearly two inches in length. "In the 
genus Hydrocharis the metasternum is prolonged somewhat, but does not 
form a long, sharp spine as in Hydrophilus and Tropisternus, and the sternum 
of the prothorax bears a keel-shaped projection. Our most common species 
is Hydrocharis obtusatus; this measures about five-eighths of an inch in 

" Some of the smaller species of this family are not aquatic, but live in 
moist earth and in the dung of cattle, where, it is said, they feed on dipterous 

The rove-beetles, Staphylinidae, form a large family, numerous in species 
and individuals over the whole country, and one whose members are readily 
recognized by the elongate flattened soft body, narrow and parallel sides, 
with short truncate leathery elytra under which the hind 
wings are compactly folded so as to be wholly concealed. 
They are mostly carrion-feeders and with the Silphidae 
(p. 261) are almost sure to be found whenever a mass of 
decaying flesh or e.xcrementitious matter exposed on the 
ground is turned over. They run swiftly when disturbed 
and curve the tip of the flexible abdomen up over the 
body in a sort of threatening way, as if they would sting. 
They cannot; they can simply smell bad. Although the 
more famihar rove-beetles are of fair size, from half an 
inch to nearly an inch long, the majority of the one 
thousand or more species found in this country — 9000 
species are known in the world — are very small. In 
California great swarms of minute rove-beetles dance in 
the air in April and May, and are a woful nuisance to 
people driving or bicycling. They get into one's eyes, 
and when crushed by rubbing, their acrid body-fluids 
both smell bad and burn. Among these smaller Sta- 
phylinids are numerous predaceous species and many which are found in 
flowers, probably feeding on pollen. Others are found on fungi, on mud, 
and in other damp places, and some live in ants' nests (see Chapter 

XV, p. 55^)- 

The larvae (Fig. 355) are found in the same places as the adults, and 
are elongate, narrow-bodied, and rather like those of the Carabidae, but 
each foot has but a single claw. The pupse of some species are enclosed 
in a sort of exudation that dries into a firm protecting coating rather like 
the horny cuticle of a lepidopterous chrysahd. 

Among the more familiar rove-beetles are species of the genus Creophilus. 

Fig. 355. — Larva 
of a rove-beetle, 
X anthaliniis 
lentus. (After 
Schiodte; twice 
natural size.) 



Fig. 356. — Rove-bee- 
tle, Creophilus vil- 
losus. (One and one- 
half times natural 

C. villosus (Fig. 356), common all over the country, is about | inch long, 

blackish, with an incomplete broad transverse patch of yellowish-gray hairs 

across the elytra and another on the second and third abdominal segments. 

Leistotrophus is a genus with but one American species, L. cingulatus, about 

same size as the preceding, but of grayish-brown color 

indistinctly spotted with brown and with a golden tinge 

on the tip of the abdomen. Staphylinus is a genus of 

twenty species or more; 5. maculosus, i inch long, is 

dark cinnamon-brown with a row of squarish black 

spots along the middle of the abdomen; S. cinna- 

mopterus, J inch long, is cinnamon-colored, with 

blackish abdomen ; S. lonientosus, \ inch long, is deep 

dull black; 5. violaceiis, \ inch long, is black with 

thorax and elytra violet. Not uncommon along sandy 

seashore in CaHfornia is a curious light-brown wing- 
less rove-beetle, Thinopimis picius, with very short 

elytra, each with an open black ring, and with a double row of small black 

dots on the abdomen. Its abdomen is short and rather broad 

Another family of carrion-beetles of comparatively few species, some of 

which, however, are familiar and widely distributed, is that of the Silphidae, 

or burying-beetles. Both adults and larvae 
feed almost exclusively on decaying flesh. 
The antennae of most species have the last 
four or five segments expanded and fused 
so as to form a conspicuous little ball or a 
compact club. Two genera include most 
o the familiar species, although the one 
hundred North American species of the 
family represent thirty different genera. 
These two are Silpha (Fig. 357), the roving 
carrion-beetles, and Necrophorus (Fig. 
358), the burying-beetles. The charac- 
teristic shape and appearance of these two 
types are well shown in the figures. The 
species of Silpha are short, broad-bodied, 
flat, dull blackish, and with the elytra rather 

leathery than horny, and lined longitudinally with shallow grooves. The 

prothorax is subcircular, with thin projecting margins. The larvce (Fig. 
359) and adults are found in and underneath putrid flesh. The larvae 
are apparently more active than the adults. Silpha lapponica, a common 
dull black form in both Europe and America, is said to enter houses in Lap- 
land to eat the stores of animal provisions. S. americana (PI. II, Fig. 5) has 

Fig. 357. Fig. 358. 

Fig. 357. — Carrion-beetle, Silpha 
noveboracensis. (One and one-half 
times natural size.) 

Fig. 358. — Burying-beetle, Necropho- 
rus marginatus. (One and one- 
half times natural size.) 



Fig. 359. — Larva 
of can ion-beetle, 
Silpha sp. (One 

the large shield-like prothorax yellowish with a black blotch in the center. 

In S. noveboracensis only the margin of the prothorax is yellow. 

The burying-beetles, Necrophorus, are large insects from an inch 

to an inch and a half long, with the body thick and parallel-sided. The 

commoner species have a pair of dull red transverse blotches on each elytron. 
In some species the prothorax and head are also marked 
with red. The common name comes from the well- 
known habit of these insects of digging underneath small 
dead animals, as mice or birds, until the corpse is in a 
hole; it is then covered over and thus really buried. 
The female lays her eggs on the corpse, and the larvae 
hatching from them feed on the decaying matter. These 
larvae have spiny plates on the back of the body and 
are otherwise unlike the Silpha larvae. Some Necrophorus 

and one-half times larvae are predaceous and others feed on decaying vege- 
natural size.) ^^^^^ j^^^^^^ 

Most of the blind, pale cave-beetles found in caves in this country and 
Europe are Silphidas. 

The Cucujidae, with a name derived from the Portuguese Cucuyo, a 
large luminous Brazilian snapping-beetle or elater, of entirely different 
family, are a family of small beetles, with flattened reddish or light-brown 
body, whose outdoors haunts are mostly under the bark of trees. Sev- 
eral species, however, have learned that 
life in a granary is just as safe from pre- 
daceous enemies, and a thousand times 
safer from starvation. Of these sophisticated 
Cucujids, Silvanus surinamensis, the saw- 
toothed grain-beetle (Fig. 360), is the most 
familiar and injurious. The adult is about 
^ inch long, flat and chocolate-brown, and 
may be distinguished from the other small 
beetles similarly attacking stored grain by 
the serrated margins of its prothorax. It 
infests dried fruits, nuts, .seeds, and dry 
pantry stores of all sorts, as well as grain bins 
and cribs. The larvae (Fig. 360) are active 
little six-legged flattened whitish grubs which run about and nibble indus- 
triously. When full-grown the larva attaches itself by a gummy excretion 
to some object, and pupates. When living in light granular substances, 
as oatmeal, etc., a delicate case is constructed of the material in which to 
pupate. In summer the life-cycle from egg to adult requires but twenty- 
four days; in spring from six to ten weeks. Six to seven generations are 

Fig. 360. — Larva, pupa, and adult of 
the saw-toothed grain-beetle, Sil- 
vanus surinamensis. (After How- 
ard and Marlatt; much enlarged.) 



produced annually in the latitude of Washington. The insect here hiber- 
nates in the adult state. 

The largest and most familiar of the outdoor Cucujids is a very flat 
bright-red species, Cucujus flavipes (PI. II, Fig. 11), about half an inch 
long, with black eyes and antennae and the legs with dark tibiae and feet. 

The Dermestidac constitute only a small family of forty or more North 
American species representing twelve genera, but one which nevertheless 
is of unusual interest and importance to entomologists, for to this family 
belong those insects which eat entomological collections. A depraved taste, 
but one which causes almost constant anxiety and occasional serious 
discouragement on the part of the industrious collector. Dermestids 
are not the bane of collectors and museum curators alone, as larder-beetles, 
"buffalo-moths," and carpet-beetles, various species of this family, help 
make life a burden to the housewife. 

All of the Dermestidae are small, oval, and plump-bodied, the largest 
species being about ^ inch long, and most of them are covered with small 
scales, which give them their rather varied colors and markings. The beetles 
themselves mostly feed on pollen, but come into houses to deposit their eggs. 
From the eggs hatch soft-bodied little grubs thickly covered with hairs, 
often very long (Figs. 361 and 362). These larvae are the real pests of house- 

FiG. 361. Fig. 362. 

Fig. 361. — Carpet-beetle or "buffalo-moth," Anthrenns scrophularia, larva and adult. 

(After Howard and Marlatt; much enlarged.) 
Fig. 362. — Black carpet-beetle, Attagenus piceits, larva and adult. (After Howard 

and Marlatt; enlarged.) 

hold and museum: they feed industriously on dried insect specimens, 
stuffed birds and mammals, woolen carpets, furs, feathers, or on meat and 
cheese (depending on the particular habits of the various species) until full- 
grown. Then they crawl into a crack or hide in the body of a museum 
specimen and pupate within the larval cuticle, which serves as a sort of thin 
hairy protecting shell. 

The usual museum pests are two species, A. varius and A. museorum, of 
the genus Anthrenus. The adult beetles are tiny, broadly oval, very convex, 
with the black body covered above with scales some of which are yellowish 



and some whitish and so arranged as to give the back an irregularly spotted 
appearance. The hairy larvae burrow into the specimens and nibble away 
at the dry bodies. Their presence may be detected by a little pile of dust 
under the pinned-up specimen and by the falling off of its legs, head, etc. 
Pour a teaspoonful of carbon bisulphide into a corner of the case and 
keep it tightly shut for a day. The fumes of the CSj are fatal to the pests. 
The carpet-beetle or " buffalo-moth " (Fig. 361) is another species, A. scrophu- 
laricE, of this same genus. The beetle is about y\ inch long, marbled black 
and white above with a central reddish line bearing short lateral offshoots 
on each side. The larva is thick, soft, active, and covered with stiff brown 
hairs. It feeds voraciously on carpets, working on the under side, and 
usually making long slits following the fioor-cracks. The beetles are common 
outdoors on plants of the family Scrophulariaceae, but come indoors to lay 
their eggs. The remedy for the carpet-beetle is to use rugs instead of 
carpets, and to lift and shake these rugs often. Another member of this 
family attacking carpets is the black carpet-beetle, Attagenus piceus (Fig. 
362). The beetle is black, and the larva is longer, more slender, and lighter 
brown than the buffalo-moth, and has a conspicuous pencil or tuft of long 
hairs at the posterior tip of the body. The larder- or bacon-beetle, Dermestes 

lardarius (Fig. 363), is about l^ inch 
long, dark brown with a pale-yellowish 
band, containing six black dots across 
the upper half of the wing-covers. 
The larva is elongate, sparsely hairy, 


Fig. 363. Fig. 364. 

Fig. 363. — The larder-beetle, Dermestes lardarius, larva, pupa, and adult. 

Howard and Marlatt; much enlarged.) 
Fig. 364. — Larva of a water-penny beetle of the Parnidce. (Four times natural 


brown, and has two short curved spines on top of the last body-segment. 
It feeds on many kinds of animal substance, as ham, bacon, old cheese, 
hoofs, horn, skin, beeswax, feathers, hair, and also attacks museum specimens. 
Another family of Clavicornia which possesses a special interest is the 
Parnidae, or "water-pennies," a family of forty species representing ten genera 
of small brown robust-bodied insects which live in water and yet do not 

Beetles 265 

have their legs fitted for swimming, nor in any other way the body partic- 
ularly modified for an aquatic life. They crawl around on submerged stones, 
sticks, and water-plants, carrying a supply of air with them, held by the 
fine pubescence of the body. The larvae are curiously flattened, broadly 
oval to nearly circular small creatures (Fig. 364), which cHng to stones and 
give the family its popular name of "water-pennies." As the legs, mouth- 
parts, eyes, etc., are all on the under side and concealed, the flat, brownish, 
leathery little "penny" is usually not recognized as an insect by the observer 
of brook life. 

The family Platypsyllidae has been estabhshed to include a single 
species of strangely shaped beetle which lives as a parasite on the bodies 
of beavers. Its name is Platypsylla castoris; it is about ^-g- inch long, blind 
and wingless, and with the elytra rudimentary. This degenerate condition 
of the body is due of course to the parasitic habit. Other obscure httle 
beetles of curious habits are the Pselaphidae and Scydmaenidse, many of 
which live commensally with ants in their nests. These beetles are rarely 
over an eighth of an inch long, and some of them have bodies strangely 
modified to look like ants. (For a further account of these insects see 
the discussion of myrmecophily in Chapter XV.) 


In this tribe of beetles, characterized by having the antennae slender, 
with each segment projecting more or less inward so as to give the whole 
antennae a saw-toothed or serrate character (Fig. 340, 10), are included sev- 
eral families certainly not closely related and having widely different habits 
and appearance. The serrate character of the antennae, too, is sometimes 
so slight that it can hardly be distinguished with certainty. The more 
important families of the tribe can b: separated by the following key: 

Head inserted in thorax as far as the eyes; body elongate or elhptical, and with unusually 
hard cuticle. 
Antennae finely serrate, the first two abdominal segments grown together on the ven- 
tral side (Metallic wood-borers.) Buprestid^e. 

Antennee often filiform; first two abdominal segments free. 

(Chck-beetles.) Elaterid^. 
Head free, but bent under the thorax. 

Small insects usually less than \ inch long (Death-watch beetles.) Pjinid.^. 

Head free, but often partly or wholly covered by the thin anterior margin of the thorax. 
Wing-covers flexible; body elongate and flattened; antennae not enlarged at tip. 

(Fireflies.) Lampyrid.e. 

Wing-covers firm, thorax convex, body not much flattened; antennae often enlarged 

at tip (Checkered beetles.) Clerid^. 

The metallic wood-borers, or flat-headed borers, a name suggested by 
the flat broad head of the larva, constitute the large and important family 



Fig. 365.— a flat- 
headed borer, 
larva of Rha- 
gium lineatum. 
(Natural size.) 

Buprestidse, of which over two hundred species occur in North America. 
The adult beedes have an elongate body, trim and compact, with a rigid 
and armor-plate-hke cuticle, and have iridescent metallic coloring. Green, 
violet, reddish, blue, copper, golden they may be, always shining like 
burnished metal and the whole body looking as if cast in bronze. The 
antennae are short and serrate on the inner margin, the head deeply inserted 
in the thorax, and the latter fitting closely against the 
abdomen and wing-covers; and the second and third 
abdominal segments are rigidly fused. These beetles are 
diurnal, running actively on tree-trunks or resting on 
flowers; seeming to delight in the warm bright sunlight, 
in which their resplendent colors flash and glance Hke 

The larvae are mostly wood-borers, although those of 
some of the smaller species mine in leaves or live in galls. 
The wood-boring Buprestid larvae are characterized by the 
strangely enlarged and flattened, legless, first thoracic 
segment, on which the small head with its powerful jaws 
sets in front, and the tapering, flattened, legless, meso- 
and meta-thoracic segments behind. The abdomen is 
elongate and rather narrow, the segments showing dis- 
tinctly. The whole larva (Fig. 365) is thus a footless whitish tadpole-like 
grub, expressively known as a flat-headed or hammer-headed borer. The 
larvs that do not burrow in wood are cylindrical and have three pairs of legs. 
The most injurious Buprestid is the notorious flat-headed apple-tree 
borer, Chrysohothris femoraia (Fig. 366), an obscure bronze or greenish- 
black beetle about half an inch long. The legs and 
under side of the body are of burnished copper, and 
the antennae green. The eggs are glued to the bark 
under scales or in cracks; the young larva on hatching 
eats inward through the bark to the sapwood and 
there burrows about, sometimes quite girdling the tree. 
Later it bores into the solid heart-wood, working up- 
ward and then again out into the bark, where it forms 
a cell in which it pupates, issuing as an adult in just 
about one year from the time of its hatching. This 
pest attacks peach- and plum-trees and several forest- 
and shade-trees as well as the apple-tree. It ranges 
over the whole country. To prevent the egg-laying on the bark, the lower 
trunk of the tree should be washed with fish-oil soap during June and July. 
When borers are once in the tree, cutting them out is the only remedy. 
The genus Agrilus contains a number of species having the head flatly 

Fig. 366. — Apple-tree 
borer, Chrysohothns 
femorata. (Twice 
natural size.) 

Beetles ■ 267 

truncate in front, as if cut sharply off, and the body rather cyhndrical than 
flattened, as with most other Buprestids. A. ruficollis, the red-necked black- 
berry-borer, y%- inch long, with dark bronze head, coppery bronze prothorax, 
and black wing-covers, has a larva that bores into the canes of blackberries and 
raspberries, burrowing spirally about in the sapwood until full-grown, when 
it bores to the pith and there pupates. The eggs are laid in June and July 
on the young canes. Infested canes often show gall-like swellings, and 
should be cut off and burned. 

Our largest Buprestids belong to the genus Chalcophora. C. virginiensis 
■ is an inch long, dark coppery or blackish with elevated lines and depressed 
spots on the elytra. The larvae bore into pines. C.liherta (PI. II, Fig. 3) is 
a beautiful pink bronze with darker raised lines. Dicerca divaricata, f inch 
long, is copper-colored, with the black-dotted elytra tapering behind and 
separated at the tips. Buprestis (PI. II, Fig. 8) is a genus of rather large 
brassy-green or brassy-black species often spotted with yellow on the elytra 
and beneath. 

Resembling the Buprestids much in general shape and appearance, the 
click-beetles, Elateridae, are readily distinguished from them by their lack 
of metaUic colors, the backward-projecting, sharp-pointed hinder angles 
of the prothorax, and their curious capacity, whence 
their name, of springing into the air with a sharp click 
when laid back downward. When a click-beetle — 
snapping-bugs and skipjacks are other common names 
for them — is disturbed it falls to the ground, lying 
there for a little while as if dead. Then if it has 
alighted, as it usually does, on its back, it suddenly 
gives a spasmodic jerk which throws it several inches 
high and brings it down right side up. This springing 
is accomplished by means of an apparatus consisting 
of a small cavity on the under side of the mesothorax ^^'^- 367- — Ventral 
into which the point of a curved projecting process click-beetle show- 
from the prosternum fits (Fig. 367). When the beetle is ing snapping appa- 
laid on its back it bends in such a way as to bring the sLe")' ^ "^a 

tip of the curved horn to the edge of the cavity, when, 
by a sudden release of muscular tension this tip slips and the insect is 
thrown into the air. The Elateridae are a large family, about 350 species 
being known in this country. They are mostly of small or medium size, 
although some are an inch or more long; a very few reach a length of 
nearly two inches. As a rule they are uniform brownish; some blackish 
or grayish and others banded and marked with brighter colors. In the 
South occur certain luminous click-beetles. In Cuba ladies sometimes use 
these phosporescent species, which are large and emit a strong greenish 



light, as ornaments, by keeping them aUve in h'ttle lace pockets on their 
gowns or attached to delicate golden chains. Two large eye-like spots on 
the prothorax, and the under side of the hinder part of the abdomen, are 
the luminous regions. 

The larvae (Fig. 368) are elongate, slender, horny-skinned, brownish 
or yellowish white, living in the ground or in decaying wood, and popularly 
and aptly known as wireworms. They have three pairs of short legs, and 
a stumpy process on the last segment of the body. They feed on the seeds, 
roots, and other underground parts of plants and do much damage to various 
crops. Often whole fields of grain are ruined by the attack of wireworms 
on the planted seeds; meadows often suffer severely, and strawberries lose 

their stolons. The beetles fly about 
in early summer, depositing their 
eggs in the ground in grassy, weedy, 
or plowed land. The larvae soon 
hatch, dig down into the soil, and feed 
on roots and seeds for two or three 
years, when they become full-grown. 
They pupate in the ground in early 
fall and the pupae transform to adults 
before winter, but the beetles do not 
issue from the ground until the fol- 
lowing spring. 

Among our largest click-beetles 
is the eyed elater, Alaus oculatus 
(Fig. 369), if inch long, blackish 
with large uneven whitish gray dots, 
a pepper-and-salt fellow, Comstock 
well calls him, with a pair of large 
white-rimmed velvet-black eye-spots 
on the prothorax. The large larvae, about 2 inches long, live in decaying 
wood and are often found in the trunks of old apple-trees. Elaier rubricollis, 
i inch long, is black with light-red prothorax; E. sangmnipennis, 
j\ inch long, is black with light-red elytra; E. nigricollis, f inch long, is 
black with whitish elytra. Athoics scapidaris, f inch long, is green sh 
black with the base of the elytra and the hind points of the prothorax clay- 
yellow. Several species of Corymbetes have the elytra brownish yellow with 
transverse zigzag black bands; C. hieroglyphicus, i inch long, has two 
bands; C. hamaius, rather smaller, has one band near the tip. Melanactes 
piceus, I inch long, is glossy black and its large larva is luminous, 
strong green light being emitted from a narrow transverse region with 
expanded ends on each segment. 

Fig. 368. Fig. 369. 

Fig. 368. — Larva of a click-beetle, Elater 

acerrimus. (After Schiodte; natural 

Fig. 369. — An eyed elater, Alans oculatus. 

(One and one-half times natural size.) 



The fireflies are familiar insects which are not flies but beetles, although 
their soft body and flexible leathery wing-covers are not of the typical 
coleopterous type. The nocturnal fireflies and their diurnal first cousins, 
the soldier-beetles, compose a coleopterous family, Lampyrida', of con- 
siderable size and common distribution over the whole world. The "glow- 
worm" of England and Europe is the wingless female of a common firefly, 
and the railway-beetle of Paraguay, a worm-hke creature 3 inches long, 
that emits a strong red light from each end of the body and a green light 
from points along the sides, is also probably the wingless female of a large 
firefly species. In this country over 200 species of Lampyridae have been 
'Ound. Comparatively few of them, however, are luminous. The light- 
giving organ is usually situated just inside of the ventral wall of the last seg- 
ments of the abdomen, and consists of a special mass of adipose tissue richly 
supplied with air-tubes (tracheae) and nerves. From a stimulus conveyed 
by these special nerves oxygen brought by the network of tracheae is released 
to unite with some substance of the adipose tissue, a slow combustion thus 
taking place. To this the light is due, and the relation of the intensity or 
amount of light to the amount of matter used up to produce it is the most 
nearly perfect known to physicists. 
Not only are the adult fireflies 
luminous, but in some species the 
pupae and larvae and even the 
eggs emit light. The combustion 
in the egg is of course accom- 
plished wholly without tracheae 
or controlling nerves. 

The larvae (Fig. 370) of Lam- 

pyrida; mostly burrow under- Fig. 370. Fig. 371. Fig. 372. 

ground, where they feed on soft- ^'?- 37o.— Larva of firefly, Photinus modestus. 
, . (Twice natural size.) 

bodied msects, slugs, and oXhtr Yig. 7,-ji.~Tue^y, Photinus scintiUans. (Three 
similar food. The adults, too, times natural size.) 

,1 r 1 r Fig. 372. — Checker-beetle, Trichodes ornatus 

are carnivorous, the diurnal forms, (Twice natural size.) 

called soldier-beetles, being com- 
monly seen on flowers or tree-trunks hunting prey. 

The commoner luminescent fireflies, or "lightning-bugs," belong to the 
genus Photinus. P. pyralis, the common species from Illinois south, is 
^ inch long, blackish, with prothorax with red disk, yellow margin, and black 
spot in center, and the elytra with narrow yellowish border. Farther north 
and east the commonest species is P. scintiUans (Fig. 371), similar in mark- 
ing but smaller. P. angulatus, h inch long, is pale, with wide yellow margins 
on elytra and the margin of the prothorax clouded with black. The com- 
moner soldier-beetles belong to the genus Chauliognathus, which is char- 



acterized by the possession of a pair of extended fleshy processes belonging 
to the maxilla, which are used in lapping up flower-nectar and pollen. Two 
common species in the East are C. pennsylv aniens, which is yellow with 
a black spot in the middle of the prothorax and one near the tip of each 
wing-cover, and C. marginatiis, which has the head and lower part of the 
thighs orange. Telephorus is another common genus without the maxillary 
processes, the species being black with the prothorax partly or wholly reddish 
yellow. The larvae of T. hilineatiis, the two-lined soldier-beetle, are velvety 
dark-brown active creatures which are very beneficial in orchards, devour- 
ing "immense numbers of such destructive beings as the larvae of the plum- 

Professor Comstock has given the name checkered beetles to the family 
Cleridae; a name apt enough for some of the species which, like the one 
shown in Fig. 372, have the body conspicuously marked with red and white 
or other colored "checks." Other species, however, content themselves 
with a monochrome coat. The family is a fairly large one, over a hundred 
species being known in this country. "The adults are found on flowers 
and on the trunks of trees running about rapidly, somewhat resembling 
brightly colored ants. Indeed some are decidedly ant-like, the prothorax 
being narrower than the wing-covers and slightly narrower than the head. 
The legs of the Clerids are rather long, the antennae with a marked knob 
at the end, and the body more or less cylindrical, either hairy or not. 

"The larvae are usually carnivorous and are most frequently found 
in the burrows of wood-boring insects, chiefly of those that live in sap-wood; 
others are found in the nests of bees, and still others feed on dead animal 
matter." The slender larvae possess short legs and a somewhat prominent 
and pointed head. They are extremely useful in keeping in check such 
destructive beetles as bark-beetles and other borers. 

The species of Clerus are prettily marked and are often found running 
about on logs and trees. C. diibins is ^ inch long, steel-blue with three 
orange bands across the elytra; C. nigrifons is j inch long, tawny yellow 
with smoky markings above and all black below; C. nigripes is similar, 
but all red below; C. sanguineus has the thorax brown and elytra scarlet. 
The species of Trichodes (Fig. 372) are hairy and prettily banded; the larvae 
live in nests of bees, and T. apiarius is a pest in beehives in Europe. 
Necrohia violacea, \ inch long or less, dark or greenish blue, is an importa- 
tion from Europe and is sometimes found in houses, but more commonly 
on carcasses and especially the bones of dead animals. It has been found 
under the wrappings of Egyptian mummies. Necrohia rnfipes, the red- 
legged ham-beetle, a red-legged steel-blue species ^ inch long, feeds on hams 
and other stored animal products. The beetles lay their eggs in May and 
June on exposed hams or other meats. The larvae hatch in a few days and 



are slender white active grubs with a brown head and brownish patches 
above and two small hooks at the end of the body. They feed on the meat 
until full-grown, when they either burrow deeper into the meat or come out 
and bore into the wooden receptacle holding it, and make a glistening paper- 
like cocoon within which they pupate. 

The family Ptinidie is composed of small obscure brownish beetles that 
would never attract our attention at all were it not for the injurious food- 
habits of many of the species. The family includes a hundred and fifty 
species, and among them a few notorious pests of rather unusual tastes. 
As the Ptinids mostly live on dead and dry vegetable matter, it was not 
improbable when I began a collecting expedition in a d ug-store that I should 
find a number of specimens of this family. But to find a majority of the 
canisters and jars containing vegetable 
drugs in the condition of roots, stems, 
leaves, etc., infested by beetles of this 
family was unexpected. The most 
abundant species on this collecting- 
ground was Sitrodrepa panicea (Fig. 

^ 7 s), which we may well call the "drug- „ ^, , , , ^• 

"^'^^' , ^■' . , ^'^ Fig. 373.— The drug-store beetle, Sitro- 

Store beetle. It was found to be drepa panicea, larva pupa, and adults, 

attacking blue-flag rhizome, comfrey- (After Howard and Marlatt; much 

, , , . , . enlarged.) 
root, dogbane-root, gmger-rhizome, 

marshmallow-root, aniseed, aconite-tuber (deadly poison to us!), musk-root, 
Indian-turnip rhizome, belladonna-root, witch-hazel leaves, powdered coffee- 
seed, wormwood stems, flowers and leaves, thorn-apple leaves, cantharides 
(dried bodies of blister-beetles), and thirty other different drugs! Larvae, 
pups, and adults were side by side in most of the canisters. Ptinus brim- 
neus, a larger Ptinid, was in half a dozen jars, and the cigarette-beetle, 
Lasiderma serricorne, suggestively named, though it feeds on tobacco in 
almost any form, was living contentedly in a jar of powdered ergot. 

"Death-watch" is a name popularly applied to several species of Ptinids 
because of their habit of rapping their heads so sharply against wood in 
which they are burrowing as to make a regular tapping or ticking sound. 
This name is claimed by species of Anobium, tiny, robust, hard-bodied, cin- 
namon-colored beetles, gV inch long, and also by Sitrodrepa panicea, our 
drug-store beetle. Comstock records finding this species breeding in 
large numbers in an old book, a copy of Dante's Divine Comedy, printed 
in 1536. Librarians wovfld call the beetle a "bookworm." 

Besides the small members of the family which feed on dried foods, 
drugs, etc., there are a few larger species of very different habits, although 
also destructive. The apple-twig borer, Amphiceriis bicaiidahis, ^ inch 
long, dark chestnut-brown above and black beneath, is the best known of 



these. It bores into live apple-twigs in early spring, entering close to a 
bud, and making a burrow several inches long for food and shelter. Twigs 
of pears and cherries are similarly infested. Both sexes bore these tunnels; 
the males have two sharp little horns on the prothorax. The eggs are laid 
in the dead or dying shoots of the greenbrier (Smilax) or in the dead shoots 
of grape. The larvaj feed on these roots or shoots and pupate in them. 
The remedy is to cut off and burn infested twigs, and to keep greenbrier 
from growing near the orchard. The red-shouldered sinoxylon, Sinoxylon 
basilare, | inch long, black with large reddish blotch at the base of each 
wing-cover, has a larva which bores into the stems of grape-vines and into 
twigs of apple and peach. This larva is a much-wrinkled grub, yellowish 
white with swollen anterior segments, three pairs of short legs, a small head, 
and an arched body. The pupa is formed inside the burrow and is of a 
pale-yellowish color. The only remedy is to remove and burn the infested 
canes and twigs. 


In this tribe are only two families, one small but containing strangely 
shaped and interesting beetles, the other very large. In both the terminal 
segments of the antennas have conspicuous lateral prolongations in the shape 
of teeth or plates (lamellas) (Fig. 340, s and 9). The families may be dis- 
tinguished as follows: 

Antennae elbowed, the club (terminal segments) composed of segments with fixed 
transverse teeth; mandibles of the male often greatly developed. 

(Stag-beetles.) Lucanid^e. 
Antennae not elbowed, the club composed of segments modified to be large flat 
plates which can be shut together like the leaves of a book; mandibles of 
males not greatly enlarged. 

(Lamellicorn leaf-chafers and scavenger-beetles.) Scarab.eid.^. 

The stag-beetles, Lucanidae, get their name from the extraordinary 
hyper-development and curious branching stag-horn-like processes of the 
males of certain of the larger, more conspicuous species. Only fourteen 
or fifteen North American species of stag-beetles are known, but the abun- 
dance and striking appearance of several of them make the family a well- 
known one. The adult beetles are found on trees, where they presumably live 
on the sap flowing from bruised places, and on honey-dew secreted by aphids 
and scale-insects. In captivity they will take moistened sugar. Comstock 
believes that some species feed on decomposing wood. The large white 
globular eggs are laid in crevices of the bark near the base of the trunk, 
and the white, soft, fat-bodied larvae (grubs) burrow into the tree either in 
rotten or sound wood, and live there for a long time. It is said that the 
larvae of some of the larger species require six years to complete their growth. 




The genus Lucanus contains four North American species, three of which 
are familiar. L. elaphiis (Fig. 374), the giant stag-beetle, of the southern 
states, varies from i^ to 2 inches in length, not including the mandibles, 
which in the male are i inch long and branched; L. dama, the common 
pinching-bug of the East, rich mahogany-brown in color, from i inch to 
I J inches long, "flies by night with a loud 
buzzy sound and is often attracted to lights 
in houses," and has a white grub larva 
looking like the white grub of the June-bug, 
but found in partially decayed trunks and 
roots of apple-, cherry-, willow-, and oak- 
trees instead of in the ground; L. placidits, 
not quite an inch long, and black, is a third 
common species. The antelope-beetle, 
Dorcus paralleliis, is less than an inch long, 
black, and with longitudinal grooves on the 
elytra. Platycercus quercus, | inch long, 
brownish black, is widely distributed. 
Ceruchus piceus, J inch long and dark brown, 
is occasionally common in rotten wood. 
The horned Passalus, P. corniitus, large 
and shining black, has a short horn bent 
forward on top of its head. 

The great family Scarabaeidae, com- 
prising over five hundred species of North 
American beetles, includes some of our most familiar kinds. Indeed 
so many common, conspicuous, and interesting Scarabaeid beetles are to be 
found by any collector, or observed by any amateur naturalist, that the two 
or three pages of this book which can be devoted to them are confessedly 
miserably inadequate to help any one. The characteristic club of the 
antenna? and heavy robust June-bug type of body make most of the members 
of this family readily recognizable. In practically all, too, the anterior 
tibiae are broad and flattened and fitted for digging. Depending on their 
habits, the Scarabaeids are readily divided into two principal groups, the 
scavengers, of which the tumble-bugs, dung-beedes, etc., are examples, 
and the leaf-chafers, of which the June-bugs, rose-bugs, rhinoceros-beetles, 
fig-eaters, and flower-beetles are examples. Some entomologists divide 
the Scarabaeids into several distinct families, but niost do not. The scavenger 
Scarabaeids are beneficial to man by their eating or burying of decaying 
matter, but the leaf-chafers are harmful, some of them being serious pests. 
The Scarabasid larvae (Fig. 376) are thick, soft-bodied, whitish, six-footed 
grubs, which usually lie curved and often on one side. They are found 

Fig. 374. — Stage-beetle, Lucanus 
elaphus male. (Natural size.) 



in manure, rotten wood, and in the ground. The familiar white grub, larva 
of the June-bug, is a typical example. 

Of the scavenger Scarabasids the tumble-bugs are wide-spread and well 
known. The species common in the East belong to three genera: Copris, 
with middle and posterior tibiae dilated at the tip; Canthon, with these tibae 
slender or only slightly dilated ; and Phaneus, with the anterior tarsi wanting, 
and the others without claws. The species of Canthon, male and female 
working together, make balls of dung, which are rolled along for some dis- 
tance and iinally buried in the ground. The female lays an egg in the ball, 

Fig. 375. 

Fig. 377. 

Fig. 376. 

Fig. 375. — Polyphylla crinita. (Natural size.) 

Fig. 376. — Larva of a large Scarabeid beetle. (Natural size.) 

Fig. 377. — Phaneus cartiifex. (One and one-half times natural size.) 

and the fat white grub hatching from it feeds on the ball until ready to pupate. 
The adult beetle issues in about two weeks from the time of laying the egg. 
The common Copris Carolina does not make a ball, but digs holes close to 
or under manure, and tills the holes with this substance, on which the larvae, 
hatched from eggs placed one in each hole, feed. The species of Phaneus 
(Fig. 377) are brilliantly colored with metallic green, rose, and bronze, and 
bear curious projecting horns on the prothorax. The famous Sacred Scara- 
beus of the Egyptians, Ateuchns sacer, was "held in high veneration by 
this ancient people. It was placed by them in the tombs with their dead; 
its picture was painted on their sarcophagi, and its image was carved in 
stones and precious gems. These sculptured beetles can be found in almost 
any collection of Eg}'ptian antiquities." 

Common dung-beetles are the numerous species of Aphodius, ^ to ^ 
inch long, with oblong, convex, or cylindrical body, and with the front 
of the head expanded shield-like over the mouth-parts. "These insects 
are very abundant in pastures in the dung of horses and cattle, and immense 
numbers of them are often seen flying through the air during warm autumn 
afternoons." Common species are yl. fimetarius, ^ inch long, with red elytra; 



A. oblongus, ^ inch long, wholly black; and A. terminalis, \ inch long, black 
with reddish legs and tips of elytra. The earth-boring dung-beetles, 
Geotrupes, have ii-segmented antennse, and the upper lip and mandibles 
can be seen from above. "The females bore holes into the earth either 
beneath dung or near it: this is to serve as food for the larvae, an egg being laid 
in each hole." G. splendidus (PI. II, Fig. 6), f inch long, dark metallic 
green to purple; G. excrementi, \ inch long, is bronze-black; G. opacus, J inch, 
is deep black. Common on dried decaying animal matter, especially skins, 
and on the hooves and hair of decaying animals are small (^ to + inch long) 
rough convex beetles, often with a crest of dirt on their elytra, belonging 
to the genus Trox. They have the thighs of the front legs greatly dilated. 
The Scaraba^id leaf-chafers are many and various in color and marking; 
they feed, when adult, on leaves, pollen, and flower-petals. They have the 
abdomen usually projecting beyond the wing-covers. The thick, fat, white, 
horny-headed larvas hve either in rotten wood or underground, feeding on 
the roots of grasses and other plants, often doing much damage in this 
way. The June-bugs or May-beetles (Fig. 378), familiar big brown or 
blackish buzzing creatures, belong to the genus 
Lachnosterna, of which sixty or more species are 
found in this country. They are but few, however, 
on the Pacific coast. The larvae are famiUar white 
grubs that live underground and feed on the roots 
of grasses, strawberries, etc. They often do much 
damage to lawns. They live as larvae 
for two or three years, and pupate in 
an underground cell. The adult 
beetles fly and feed at night, often 
injuring the fohage of cherry, plum, 
and other trees. The familiar rose- 
chafer, Macrodactylus siibspinosus 
(Fig. 379), I inch long, a slender 
Pig. :;y8. Fig. 37Q. yellowish beetle with pale red legs, 

Fig. 378. — The June-beetle, Lachnosterna does great damage to roses and grapes, 
fusca. (One and one-half times natural ^^ppgaring in early summer and eat- 

FiG. 379.— The rose-beetle, Macrodactylus ing flowers and foliage. The larvae 
subspinosus. (Twice natural size.) jj^.g underground, feeding on the roots 

of various plants, but especially grasses. The spotted vine-chafer, Pelidnota 
punctata (PI. II, Fig. i^), i inch long, stout, convex, polished reddish or 
yellowish brown, with three large black dots on each elytron, with under 
side of body metallic greenish black, flies during July and August by day, 
feeding on grape-leaves. The larva lives in rotten wood, especially the 
decaying roots of apple, pear, hickory, and other trees. It pupates in a 



cell in the wood. The goldsmith-beetle, Cotalpa lanigera, of similar size 
and shape, is gUstening, burnished lemon yellow above with metallic 
greenish, golden, and rose reflections; below it is copper-colored and 
thickly covered with whitish wool, hence the name lanigera, or wool-bearer. 
It appears in May and June, flies by night, and feeds on the foliage of 
various trees. The larva lives in the ground, feeding on plant-roots. It is 
said to require three years to complete its growth. 

The largest beetles in our country are the oddly shaped rhinoceros-beetles, 
Dynastes, found in the south and west. D. titynis (Fig. 380), 2^ inches long, 
is greenish gray with scattered black spots on the elytra; the male has a 
large horn on the head and three horns, one larger than the others, on the 
prothorax; the female has only a tubercle on the head; it is a southern species. 
D. grantii, of the west, has the large prothoracic horn twice as long as in 
tityrus. In the West Indies occurs D. hercules, six inches long! The larvae 

Fig. 380. — The rhinoceros-beetle, Dynastes tityrus. (Natural size.") 

(Fig. 381) of these beetles live in the roots of decaying trees. Allied to 
Dynastes is the genus Ligyrus, of which L. rugiceps, the black sugar-cane 
beetle of the southern states, is the best -known species; it burrows into the 
base of sugar-cane and sometimes corn, and is often seriously destructive. 
The larva lives in manure. The flower-beetles are Scarabseids of several 
genera, which are commonly seen flying from flower to flower and feeding 
on pollen. The bumble flower-beetle, or Indian Cetonia, Euphoria inda 
(Fig. 382), a common species, is | inch long, yellowish brown, with the 
elytra irregularly covered with small blackish spots, and with the whole 
body clothed with short fox-colored hairs, it appears early in spring, and 
flies near the ground with a loud humming. It feeds on flower-pollen, the 
tassels and green silk of young corn, and later on ripening fruits of all kinds; 
it often swarms about wounded trees, lapping up the escaping sap. The 
larvae feed on decaying substances underground. The fig-eater, or "southern 
June-beetle," AUorhina nitida, f inch to i inch long, is rather pointed in 



front, velvety green with the sides of thorax and head brownish yellow; the 
under side is not velvety, but metallic green. It flies with a loud buzzing 
sound and feeds on ripe fruit. The larva; are found in richly manured 
soil, feeding on decaying matter. They cannot use the short legs for crawling, 
but move along on their backs by means of stiff bristles. "If put on a table 

Fig. 381. Fig. 382 

Fig. 381. — Larva and pupa of the rhinoceros-beetle, Dynasius tityrus. (After Chittenden; 

one-half natural size.) 
Fig. 382. — Euphoria inda. (One and one-half times natural size.) 

in normal position, they immediately turn upon their backs and by the 
alternate contractions and expansions of their body-segments they wriggle 
away in a straight line." 


In the four families of beetles constituting this- section the feet are appar- 
ently composed of four tarsal segments, one of the more usual five being 
so reduced in size and fused with the last segment as to be practically indis- 
tinguishable as a distinct segment (except in the Spondylidae) . The first 
three tarsal segments are dilated and furnished with brushes of hairs on 
the sole, the third segment being plainly bilobed (Fig. 341, 12). This 
section is sometimes named Phytophaga, because of the voracious plant- 
feeding habits of all its members. The three principal famihes of the 
section can be separated by the following key: 

Body short and more or less oval; antennae short. 

Front of head not prolonged as a short broad beak; elytra usually covering the tip 
of the abdomen; both larvce and adults live on green plants. 

(Leaf-beetles.) Chrysomelid^. 
Front of head prolonged as a short, quadrate beak; elytra rather short, so that the 
tip of the abdomen is always exposed; larvae live in seeds. 

(Pea- and bean-w^eevils.) Bruchid.e. 

Body elongate; antennae almost always long, often longer than the body; larvae are 

wood -borers (Long-horn beetles.) Cerambycid.^. 

The leaf-beetles, Chrysomelidoe, are one of the largest of the beetle fami- 
lies, over 600 North American species being known. They are mostly small, 

278 Beetles 

the familiar Colorado potato-beetle being one of the largest species in the 

family; the body is short, more or less oval in outUne, strongly convex above; 

the head small, much narrower than the prothorax, and with the antennae 

inserted widely apart. The adults walk slowly about on the plants on which 

they feed, and when disturbed usually fold up the legs and fall, inert, to the 

ground. However, they sometimes take readily to wing. The eggs are 

usually laid in little groups on the food-plants, and the larvae, rather broad, 

thick, and roughened, crawl about, exposed, on the leaves which they eat. 

Sometimes they eat only the soft tissue of the leaf, skeletonizing it; some mine 

inside the leaf, and a few burrow into- stems. Most, however, eat ragged 

holes in the leaves, and, if feeding on cultivated plants, do great injury. 

Indeed there are perhaps more beetle enemies of our crops, shade-trees, and 

ornamental plants in this family than in any other in the order. 

The Colorado potato-beetle, Doryphora lo-lineata (Fig. 383), with 

robust, oval, cream-colored body, and elytra with five longitudinal black 

stripes on each, is a notorious Chrysomelid whose gradual extension or 

migration eastward from its native home in Colorado 

created much excitement forty years ago. Its native 

food-plant is the sand-bur, Solanum rostratum, a 

congener of the potato, but after 1850 it began to find 

its way to the potato-plants of the early settlers; by 

1859 it had reached Nebraska, 1861 Iowa, in 1864 

and 1865 it crossed the Mississippi and gradually 

Fig. 383. — The Colo- gxtehded eastward until 1874, when it reached the 
rado potato - beetle, . , . ^ 
Doryphora lo-lineata. Atlantic Ocean. Fmally It obtained a partial foothold 

(Twice natural size.) jn Europe, creating great consternation there, but it has 
never got to be a serious pest across the ocean. The orange-red eggs are 
laid on the leaves, and the larvae are curious humpbacked soft-bodied crea- 
tures with black head and Venetian-red body. They crawl down and bur- 
row into the ground to pupate. There are three generations a year in the 
latitude of St. Louis, the beetles of the last brood crawling underground 
to hibernate. 

The common asparagus-beetle, Crioceris asparagi, red, yellow, and black, 
gnaws holes in young asparagus-heads, and the brown slug-Hke larvae which 
hatch from oval blackish eggs laid on the heads also eat them. The three- 
lined Lema, Lema trilineata, of similar shape, but yellow with three longi- 
tudinal black stripes on each elytron, is common on "ground-cherries." 
Their larvae have the curious habit of covering their backs with their own 
excrement. Elm-trees in the East are often badly infested with the imported 
elm-leaf beetle, Galerucella luteola (Fig. 384), a common European pest. 
It first got to this country in 1834 and is now "in all probability responsible 
for more ruined elm-trees in the Hudson River valley than all other destruc- 



tive agencies combined." The beetle, ^ inch long, is reddish yellow with 
black spots on head and prothorax, and a thick black stripe on each elytron. 
From orange-yellow eggs laid on the under side of the leaves hatch larvae 
which when full grown are ^ inch long, flattened, marked with blackish 
and yellow. They skeletonize the leaves. When ready to pupate they 
crawl down into the ground. The beetles themselves after issuance fly back 
to the tree-tops and eat holes in the leaves. There are two broods a year, 
and the adult beetles of the last brood hibernate in concealed places. 

Fig. 384. — The elm-leaf beetle, Galerucella liiteola; eggs, larvae, pupa, and adults. (After 
Felt; eggs greatly magnified; larvae, pupa, and adults about twice natural size.) 

Four species of the genus Diabrotica are common over the country and 
very injurious: D. vittata, the striped cucumber-beetle, is greenish yellow 
with two black stripes on each elytron, and feeds on cucumber-, pumpkin-, 
squash-, and melon-vines, the larva also burrowing into the stems and roots 
of the same plants; D. 12-piinctata (Fig. 385) is greenish yellow with six 
black spots on each elytron, and feeds on a great variety of plants, the larva 
often being injurious to corn in the South; D. longicornis, the corn-root- 
worm beetle, is grass-green with spots or stripes, and its underground larva 
is very destructive to corn by burrowing into its roots; D. soror (Fig. 386)^ 
of the Pacific coast, the flower-beetle or "diabrotica," yellowish green with 



six black spots on the wing-covers (like l2-punctata), does great damage as 
an adult by eating into the flower-buds of roses, chrysanthemums, and a 
host of others, the larva feeding on the roots of alfalfa, chrysanthemums, 
and many other plants. 

Fig. 385. Fig. 386. 

Fig. 385. — The cucumber-beetle, Diabrotica 12-punctata. 
Fig. 386.— The California flower-beetle, Diabrotica soror. 
Fig. 387. — Chrysomela digsbyana. (Twice natural size.) 

Fig. 387. 

(Three times natural size.) 
(Three times natural size.) 

Chrysochus auratus (PI. II, Fig. 4), f inch long, golden green in color, 
found in the East, and C cobaltinus (PI. II, Fig. 7), of same size and shape, 
but brilliant blue, found in the West, are the two most beautiful Chrysomelids. 

Chrysomela (Fig. 387) is a 
genus whose species are often 
curiously marked with short, 
curved lines and irregular 
spots. The active little flea- 
beetles, with swollen hind 
femora, and able to leap vigor- 

Fig. 388. Fig. 389. 

Fig. 388.— Larvae of the grape-vine flea-beetle, Haltica chalybea. (After Slingerland; 

much enlarged.) 
Fig. 389.— a tortoise-beetle, Coptocyda aurichalcea. (Two and one-half times natural 


ously, are common pests of grapes, cucumbers, melons, cabbages, turnips, etc., 
numerous species being known. They are small, usually about -^\ to \ inch 
long, and commonly blackish or steel-blue in color. Haltica chalybea, the steel- 
blue flea-beetle (Fig. 388), is common on grape-vines, where it feeds on the 



fruit and leaves; Crepidodera aicimieris, the cucumber flea-beetle, ^V i^^ch 
long, and black, attacks melons, cucumbers, and other vegetables. The 
tortoise-beetles (Fig. 389) are curiously shaped, flat beloM^, convex above, and 
with the prothorax and elytra thinly margined so as to give them a tortoise-like 
appearance from above; they are usually iridescent greenish and golden in color, 
and are often called goldbugs. The colors appear and disappear strangely 
while the insects are alive, but are always lacking in the dead specimen. 
Coptocycla clavata has two projections of the central dark color of each 
elytron looking like the four short broad legs of a tortoise; Cassida bicolor 
is like "a drop of burnished gold"; Chelymorpha argiis, | inch long, brick- 
red with many black spots on prothorax and elytra, is found on milkweeds; 
Physonota imipunctala, ^ inch long, the largest of our tortoise-beetles, yellow 
with whitish margins, is common in midsummer on wild sunflowers. 

The small familv Bruchidae contains two common and important beetles, 
viz., the pea-weevil, Bntchus pisl (Fig. 390), and the bean-weevil, B. 
obtectiis (Fig. 391). The adult pea-weevil is \ inch long, general color rusty 
or grayish black with a small white spot on the thorax. The eggs are small, 
fusiform, and yellow. The grubs on hatching bore through the pod into 
the peas. The hole made in the growing pea soon closes up, leaving 
the voracious larva within. Here it often comes to an untimely end, 
— which is uncomfortable to think about. If, however, the peas are 
allowed to ripen and are put away for seed, it eats on until there is 

Fig. 3QO. Fig. 391. 

Fig. 390. — The pea-weevil, Bnichus pisi, and an infested pea. (Natural size of beetle 

indicated by line.) 
Fig. 391. — The bean-weevil, Bnichus ohtectus, and an infested bean. (Natural size 

of beetle indicated by line.) 

only a shell left of the pea. Weeviled peas are unfit for food, and, as 
proved by the experiments of Professor Popenoe, should not be used for 
seed. During the fall and winter the larvae pupate and finally mature as 
weevils (the adult beetles). Some of the beetles emerge from the peas, 
while others remain in them until they are planted. 



"Weevily" peas should be put into a tight box or bin, together with a 
small dish of bisulphide of carbon, the fumes of which will kill the insects. 
Or they may be immersed for a minute or two in water heated to 140'=' F.; 
this will kill all the beetles and larvas. 

The bean-weevil is a little larger than the pea-weevil and lacks the 
white spot on the thorax. Its Hfe-history is about the same as that of the 
pea-weevil, the eggs being laid of course on the young bean-pods. Several 
eggs are frequently laid in a single bean. The bean- weevil continues to 
breed also in dry stored beans, and increases its damage materially if the 
stored beans lie long untouched. It is therefore necessary to treat weeviled 
beans with bisulphide of carbon or hot water before storing them away. 

Fig. 393. 

Fig. 392. — Prionus californiciis. (Natural size.) 

Fig. 393. — Larva of Ergates spiculatiis. (Natural size.) 

Fig. 392. 
392. — Prionus californiciis. 

The other principal tetramerous family besides the Chrysomelids is the 
Cerambycidae, or family of long-horn wood-boring beetles: "long horn" 
because of their long slender antennae, and "wood-boring" because their 
larvffi live in burrows in the trunks of trees. The beetles themselves are 
usually large and strikingly colored and patterned, and whenever seen 
attract attention. Nearly 600 species are known in North America, and 
they are common all over the country. As might be concluded from the 
habits of the larvae, the family includes numerous serious pests, such species 
as the round-headed apple-tree borer, the oak-pruners, various hickory- 
borers, the twig-girdlers, the giant Prionids et al., all causing much damage 
to orchards and forests. 




The eggs are usually laid on the bark, and the whitish, usually footless 
soft-bodied but hard-headed and strong-jawed larvae burrow about in the 
tree-trunk for a year or two or even three (varying with the different species) 
feeding on the chewed wood. They pupate in the burrow, in a cell par- 
titioned off with chips, or sometimes specially made just under the bark 
The beetle has only to gnaw its way through the bark or the loosely plugged 
burrow to escape from the tree. These wood-borers usually select a 
weakened or dying tree for attack. 

The largest Cerambycids belong to the subfamily Prionida? (Fig SQ2) 
whose members have the sides of the prothorax sharply margined and 
usually toothed. Prionus laticoUis, the broad-necked Prionus, varies from 

Fig. 394.— The sugar-maple borer, Plagionotus speciosiis, larva; and adult beetle. (After 

Felt; natural size.) 

I inch to 2 inches in length, and is pitchy black or brown, the prothorax 
with three sharp teeth on each lateral margin, and the antennae 12-segmented; 
the larvffi, which live three years, are great footless white grubs, 2^ to 3 
inches long, which burrow in the roots of oak, poplar, cherry, apple 
grape-vine, and blackberries. The tile-horned Prionus, P. imbricornis 
a similar beetle, has nineteen antennal segments in the male and usually 
Sixteen in the female; OrtJiosoma brunnea, is long (li to 2* inches) and 



narrow, with the margins of the body nearly parallel. In the south occurs 
the genus Mallodon, and on the Pacific coast the genus Ergates (with a 
single species, spiailatHs), both 2 J inches long, and with the lateral margins 
of the prothorax with many fine sharp teeth. The larvae (Fig. 393) of 
Ergates live in the giant sugar and yellow pines of the Sierra Nevada forests. 
The cloaked knotty-horn, Desmocerus palliatus (PI. II, Fig. i), is a 
beautiful species, dark greenish blue with the bases of the elytra orange- 
yellow; the larvae bore in elder-pith. Cyllene robinicr, the locust-borer (PI. II, 

Fig. 395. 

-Maple-tree borer, Elaphidion villosum, larva, pupa, and adult beetle, 
(After Felt; natural size.) 

Fig. 15), is black, with striking yellow bands often found on goldenrod; 
its larvae live in locust-trees. A similar species, Cyllene pictus, attacks the 
hickory. The red milkweed-beetle, Tetraopes tetraopthalmus (PI. II, 
Fig. 10), brick-red with black spots, is a common species on milkweeds; 
the larvae bore into the lower stems and roots. Two beautiful Cerambycids 
of California are shown in Figs. 2 and 1 6 of PI. II. 

The sugar-maple borer, Plagionotiis speciosus (Fig. 394), is a serious 
pest of sugar-maples in New York and elsewhere in the East. The beetle, 
I inch long, is black, brilliantly marked with yellow; the eggs are laid in 

Beetles 28 c 

July or August in the bark, the young borer (a footless, flattened, whitish 
grub) burrowing first into the sap-wood, where it passes the winter. Dur- 
ing the next year it bores vigorously around under the bark, and when about 
sixteen months old makes a final deep burrow into the heart-wood, in the 
end of which it pupates. Fig. 394 shows all the stages of this insect. The 
maple-tree pruner, Elaphidion villosum (Fig. 395), f inch long, slender 
grayish brown, lays its eggs on small twigs in maple-trees in July; the larvae 
bore into the center of the twig, eat out a large portion of the woody fiber, 
plug the end of the burrow with castings, and wait for a strong wind to break 
off the nearly severed branch. In the fallen twigs thus broken off the 
larvae pupates, and the beetles issue, the life-history taking just about a year 
for completion. This pest also "prunes" oaks, and apple, pear, plum, and 
other fruit trees. The sawyers, various species of the genus Monohammus, 
are beautiful brown and grayish beetles with extremely long delicate antennc-e; 
the larvae bore in sound pines and firs and do great injury to evergreen 

One of the worst and most famihar orchard pests is the round-headed 
apple-tree borer, Saperda Candida (Fig. 396). The beetle is f inch long, 
narrow, and subcylindrical, pale brown with 
two broad creamy-white longitudinal stripes. 
The eggs are laid on the bark at the base of 
the tree in June and July. The larva works 
at first in the sap-wood, making a flat shallow 
cavity filled with sawdust and castings; later 
it burrows deeper and works upward. When 
nearly three years old it bores a tunnel from 
the heart-wood out nearly to the bark, partly F,o^3Q6.-The round -heLd 

fallmg the outer part with sawdust and then apple-tree borer, Saperda cau- 

retires to the inner end and pupates. Two f?/' ^^^^"^ .''"'^ '"^''^^ ^^'^t'^- 

.1 1 r. . , . , (After Saunders; natural size.) 

or three weeks after pupation the adult beetle 

issues from the pupal skin, works outward along the tunnel and cuts a 
smooth circular hole in the bark through which it escapes. When several 
larvas are working in a tree they may completely girdle it, so that it dies. 
The most effective remedy is to apply a repellent wash of lime or soft soap 
from the base of the trunk up to the first branches several times during the 
egg-laying time, i.e., June and July. 

A small family, Spondylida?, called the aberrant long-horned beetles, is 
represented in North America by four species, of which the most common 
is Parandra brunnea (PI. II, Fig. 14), a beautiful polished mahoganv- 
brown beetle found under the bark of pine-trees. 

286 Beetles 


Only one family is included in this section of beetles with but three tarsal 
segments in each foot, namely, the familiar, little ladybirds or plant-louse 
beetles, the Coccinellidte. Their uniformly small size, the semispherical shape, 
and the "polka-dot" pattern distinguish them readily from all other beetles 
except perhaps the Chrysomelids, a few of which are often mistakenly 
called ladybirds. This is a particularly unfortunate confusion because of 
the radically different food-habits and consequent economic relation to 
man of the two families. The Chrysomehdae, or leaf-eaters, both as larvse 
and adults, attack our crops and trees and flowers; the CoccinelHdae, or 
ladybirds, both as larvae and adults, feed on plant-lice and scale-insects, 
great enemies of our orchards and gardens, and thus are among our best 
insect friends. A friend of mine found that his roses were suffering from 
insect attack; he saw httle, convex, black-spotted reddish beetles clamber- 
ing busily up and down the stems, and he set to work to pick these off one 
by one and drop into a tin cup with petroleum in the bottom. When he had 

# i ^ $ 

Fig. 397. — Some Californian ladybird -beetles; beginning at left of upper row the species 
are Megilla vitigera, CoccineUa caJijornica, C. ocidata, Hippodamia convergens; 
beginning at left of lower row, CoccineUa trifasciata, C. sanguinea, C. ahdoniinalis, 
Megilla maculata. (Twice natural size.) 

a full pint he showed them proudly. But the more little round beetles he 
picked off the more rapidly wilted his roses. And for the wholly sufficient 
reason that he was collecting and killing the ladybirds that were making 
a fight — a losing one in the face of my friend's active part in it — against 
the hosts of tiny inconspicuous green rose-aphids that were sucking the sap 
out of the rose-stems and buds. So be it remembered that not all bugs 
are bad bugs, but that some, like the ladybirds, are most effective helpers 
in waging war against the real pests! 

There are about 150 species of ladybirds known in the United States^ 
and almost all are reddish brown with black dots or black with reddish 

Beetles 287 

spots. Their colors and markings make them conspicuous, and vet the 
natural enemies of insects, the birds, obviously let them alone; it is presumed, 
therefore, that these beetles are ill-tasting to birds, and that their bright colors 
are of the nature of readily perceived warning signs (see discussion of 
this subject in Chapter XVII). 

The eggs are laid on the bark, stems, or leaves of the tree or plant on 
which aphids or scale-insects are present. Sometimes they are deposited 
in little patches right in the middle of a colony of plant-lice. The larva? 
(Fig. 398) are elongate, widest across the prothorax and tapering back to 
the tip, with the skin usually roughened or punctate, bearing hairs and short 
spines, and marked with blackish, reddish, and yellowish. The larvae feed 
steadily on the soft defenceless aphids or young scale-insects, or on the eggs 
and young of other larger insects. When full-grown they pupate, attached 
to the leaves or stems without entirely casting off the last larval exuvia (Fig. 
398). This cuticle often surrounds the pupa "like a tight-fitting overcoat 
with the front not closed by buttons." In other cases the larval skin is 
forced backwards and remains as a little crumpled pad about the posterior 

The two-spotted ladybug, Adalia bi punctata, reddish yellow with a 

single black spot on each elytron, is common in the East, where it often 

enters houses to hibernate. The nine-spotted 

ladybird, Coccinella novemnotata, has yellowish 

elytra with four black spots on each in addition 

to a common spot just behind the thorax. 

The "twice-stabbed" ladybird, Chilocorus 

bivulnerits, is shining black with a large red 

spot on each elytron. Anatis ij-punctata, the 

fifteen-spotted ladybird, is a large species with 

dark brownish-red elytra bearing seven black 

spots each, and a median common spot just 

behind the thorax. i?,^ • o ^ 1 j i,- j u .1 

Fig. 398. — A ladvbird-beetle, 

In California the ladybirds are of great Coccinella caliloriiica ; larva, 

importance to the fruit-growers, their steadv P'^''''^' ^"'^.^'^1'^ °'' Lawson's 
, , , , . r 1 • , . ' cvprees. (Twice natural size.) 

wholesale destruction of scale-msects bemg an 

important factor in successful fruit-raising. Fig. 397 illustrates eight species 
found on the Pacific coast. A number of ladybird species have been imported 
from Australia and other countries from which numerous destructive scale- 
insects had been earlier unwittingly brought on nursery stock. Most conspic- 
uously successful of these attempts to introduce and disseminate original home 
enemies of imported pests has been the estabhshment of the small red-and- 
black ladybird, Vedalia cardinalis, which feeds exclusively on the fluted or cot- 
tony cushion-scale (Icerya purchasi) (Fig. 254). This Australian scale first 

288 Beetles 

appeared in California near Menlo Park in .1868 on orange-trees, and in a 
few years had become so abundant and widely spread over the state that 
it seriously threatened the extinction of the great orange industry. In 1888 
a few live Vedalias (altogether about 500 specimens in five separate lots) 
were brought from Australia, put on trees infested by the fluted scale, and 
by helpful scattering of the progeny of these original emigrants this lady- 
bird species was soon distributed to all scale-infested localities. In a few 
years it had the pest completely under control, and has ever since remained 
its master. And California continues to grow Washington oranges. 


This section includes those beetles which have the front and middle 
feet with five tarsal segments, the hind feet with four. It is a heterogeneous 
assemblage, including, besides two large families of u'idely differing aspect 
and habits, a number of small ones of obscure, little known, and mostly 
uncommon species of small size, which present a wide variety of structure 
and life-history. The two principal families can be distinguished by the 
following diagnosis: 

Head without distinct neck, narrower than thorax and more or less inserted in it; 
body-wall hard; color usually black. 

(Darkling ground-beetles.) Tenebrionid^. 

Head as wide as prothorax, and attached to it by a visible neck; body soft and 

elytra flexible; colors often diversified, frequently metallic blue or green 

(Blister- and oil-beetles.) Meloid.c 

The common ground-beetles of the North and East are the swift preda- 
ceous Carabidae; any stone or log turned over 
will reveal them. In the dry warm western plains 
and southwestern semi-desert states, however, the 
slower vegetable-feeding Tenebrionidae are the com- 
mon ground-beetles. The most familiar of them on 
the Pacific coast are large, awkwardly moving, shin- 
ing black pinacate bugs, Eleodes (Fig. 399) which, 
when disturbed by the turning over of their covering 
stone, stand on their fore legs and head and emit an 
ill - smelling fluid from the tip of the abdomen. 

Yi(y ,gQ_ Pinacate bug, They have no wings, and the thick horny elytra are 

Eleodes sp. (Natural grown fast to the back. All the rest of the body 
^^^^■•^ is similarly armor-plated, and the collector has to use 

an awl to make a hole through the body-wall for pinning up his specimens. 

Beetles 289 

The darkling-beetles constitute a large family, more than four hundred species 
being known in this country, although comparatively few of them are at 
all familiar. They are mostly dull or shining black, and feed on dry vege- 
table matter, often in a state of decay. Some live in grain, flour, meal, or 
sawdust; others in Hving or dead fungi, and a few are probably predaceous. 
A common species in mills, stables, grocery-stores, and pantries is the meal- 
worm beetle, Tenehrio nwlitor, ^ to f inch long, flattened, brownish, with 
squarish prothorax and longitudinally ridged elytra. The stout, cylindrical, 
hard-skinned, waxy, yellowish-brown larvae, or meal-worms, infest flour 
and meal. They are often bred by bird-fanciers as winter food for insect- 
eating song-birds. For this purpose they are raised in large numbers in 
warm boxes partly filled with bran, in which they undergo all their metamor- 
phosis. T. obscurus is a darker, almost black, species found also in mills 
and granaries. Both of these species have been spread all over the world 
by commerce. A smafler brown species, Echoceriis maxillosiis, ^ inch long, 
is common in the southern states in old and neglected flour. 

Uloma impressa, J inch long, deep mahogany-brown, is common in the 
east, occurring in decaying logs and stumps. Smaller species of the same 
genus, lighter in color, are also to be found in 
similar places. An odd-looking species called 
by Comstock the forked fungus-beetle, Boleto- 

therus bijurcus, is not uncommon in the north Fig. 400. Larva of a Tene- 

and east in and about the large shelf-fungi brionid, Boletotherus bifurcus. 
/r> 1 \ iU i " iU -J r i (Twice natural size.) 

(rolyporus) that grow on the sides of trees. 

The surface of the body and elytra is very rough, and two conspicuous 
knobbed horns project forward from the prothorax. The larva; (Fig. 400) 
live in the fungi. 

The other of the two larger heteromerous families, the Meloidae, numbering 
about 200 North American species, includes beetles of unusual structural 
character and appearance, of peculiar physiological properties, and of a 
highly specialized and unique kind of metamorphosis. The Meloids are 
known as oil-beetles from the curious oily fluid emitted by many species 
when disturbed, and as blister-beetles from the inflammatory and blistering 
effect of the application of the pulverized dry body svibstance to the human 
skin. This powdered blister-beetle is known to pharmacists as cantharides, 
and is a recognized therapeutic substance. The beetles are rather long 
and slender-legged and have a soft fleshy body with flexible wing-covers 
which are sometimes rudimentary, being then short and diverging (Fig. 
401). The head is broad and set on a conspicuous neck, and hangs with 
mouth downward. They are to be found crawling slowly about over field- 
flowers, as goldenrod, buttercups, etc., often in companies of a score or more 
individuals. Many of the species are brightly colored, metallic bronze, 



green, blue, and steel-black being common colors (PL II, Fig. 12). Some, 
however, are grayish, dead black, or yellowish and brown. All are leaf-feeders- 
In the development of the blister-beetles an extreme condition known 
as hypermetamorphosis occurs, which is undoubtedly the result of a purpose- 
ful adaptation brought about by long selection, but 
which seems an almost impossible achievement of 
such "blind" natural forces. The eggs are deposited 
in the ground; from them hatch minute active strong- 
jawed larvae (Fig. 402) with three pairs of long legs, 
each terminating in three claw-like spines. These 
larvse are called triungulins. They run about 
seeking food, which, varying with different species, 
consists of the eggs of locusts, or the eggs and 
honey of solitary bees. The triunguhn of Epicauta 
Fig. 401. — The striped vittata, one of our common Meloid species, studied 
potato-beetle, Epicauta ]^y Riley, explores cracks and burrows in the ground 
^wLTnatimrsize^r'''' until an egg-pod of a locust (usually of one of the 
destructive Melanoplus species) is found. Into this 
the triungulin burrows and begins to devour the eggs. After a few days 
given to eating a couple of eggs it moults and appears in a very different 

Fig. 402. — Hypermetamorphosis of Epicauta vittata. A, young larva or triungulin; 
B, caraboid larva; C, coarctate larva; D, scarabasoid larva; E, pupa; F, adult. 
(After Riley; natural size indicated by line.) 

larval guise with soft skin, short legs, small eyes, and different body form 
and proportions. One week later a second moult occurs, but without re- 

Beetles 291 

vealing much of a change in the larva, although it is now more curved, less 
active, and somewhat like a small June-beetle grub; after a third moult it is 
still more helpless and grub-like. It now grows rapidly. When full-grown 
it leaves the ruined egg-pod, makes a little cell in the ground near by in 
which it lies motionless except for a gradual contracting and slow fourth 
moulting, after which it appears as a completely helpless semi-pupa, or 
coarctate larva. In this state it passes the winter. In spring the fifth 
moult takes place, leaving the larva much as before, only smaller and 
whiter. It becomes now rather active and burrows about, but takes no 
food, and after a few days again moults for the sixth time, to appear at last 
as a true pupa. Five or six days later the adult beetle emerges. 

Those blister-beetles which live parasitically on bees' eggs instead of on 
those of the locust probably follow about the course described by Fabre 
for Sitaris humeralis, a European species, an account of which I quote 
from Sharp (Cambridge Natural History, vol. vi): "The eggs of the Sitaris 
are deposited in the earth in close proximity to the entrances to the bees' 
nests, about August. They are very numerous, a single female producing, 
it is believed, upward of two thousand eggs. In about a month — towards 
the end of September — they hatch, producing a tiny triungulin of black 
color; the larvas do not, however, move away, but, without taking any food, 
hibernate in a heap, remaining in this state till the following April or May, 
when they become active. Although they are close to the abodes of the 
bees, they do not enter them, but seek to attach themselves to any hairy object 
that may come near them, and thus a certain number of them get on to the 
bodies of the Anthophora [the bees] and are carried to its nest. They 
attach themselves with equal readiness to any other hairy insect, and it is 
probable that very large numbers perish in consequence of attaching them- 
selves to the wrong insects. The bee in question is a species that nests in 
the ground and forms cells, in each of which it places honey and lays an 
egg, finally closing the receptacle. It is worthy of remark that in the case 
of the Anthophora observed by M. Fabre the male appears about a month 
before the female, and it is probable that the vast majority of the predatory 
larva2 attach themselves to the male, but afterwards seize a favorable oppor- 
tunity, transfer themselves to the female, and so get carried to the cells of 
the bee. When she deposits an egg on the honey, the triungulin glides from 
the body of the bee on to the egg, and remains perched thereon as on a raft, 
floating on the honey, and is then shut in by the bee closing the cell. This 
remarkable act of slipping on to the egg cannot be actually witnessed, but 
the experiments and observations of the French naturalist leave little room 
for doubt as to the matter really happening in the way described. The egg 
of the bee forms the first nutriment of the tiny triungulin, which spends 
about eight days in consuming its contents; never quitting it, because con- 

292 Beetles 

tact with the surrounding honey is death to the httle creature, which is 
entirely unfitted for hving thereon. After this the triunguhn undergoes 
a mouh and appears as a very different creature, being now a sort of 
vesicle with the spiracles placed near the upper part; so that it is admirably 
fitted for floating on the honey. In about forty days, that is, towards the 
middle of July, the honey is consumed, and the vesicular larva after a few 
days of repose changes to a pseudo-pupa within the larval skin. After 
remaining in this state for about a month some of the specimens go through 
the subsequent changes, and appear as perfect insects in August or Septem- 
ber. The majority delay this subsequent metamorphosis till the following 
spring, wintering as pseudo-pupai and continuing the series of changes in 
June of the following year; at that time the pseudo-pupa returns to a larval 
form, dififering comparatively little from the second stage. The skin, 
though detached, is again not shed, so that this ultimate larva is enclosed 
in two dead skins; in this curious envelope it turns round, and in a couple 
of days, having thus reversed its position, becomes lethargic and changes 
to the true pupa, and in about a month subsequent to this appears as a 
perfect insect, at about the same time of the year as it would have done 
had only one year, instead of two, been occupied by its metamorphosis. 
M. Fabre employs the term third larva for the stage designated by Riley 
Scolytoid larva, but this is clearly an inconvenient mode of naming the stage. 
. . . Meloe is also dependent on Anthophora, and its life-history seems 
on the whole to be similar to that of Sitaris; the eggs are, however, not 
necessarily deposited in the neighborhood of the bees' nests, and the 
triungulins distribute themselves on all sorts of unsuitable insects, so that 
it is possible that not more than one in a thousand succeeds in getting access 
to the Anthophora nest. It would be supposed that it would be a much 
better course for these bee-frequenting triungulins to act like those of Epicauta, 
and hunt for the prey they are to live on; but it must be remembered that 
they cannot live on honey; the one tiny egg is their object, and this appar- 
ently can only be reached by the method indicated by Fabre. The history 
of these insects certainly forms a most remarkably instructive chapter in 
the department of animal instinct, and it is a matter for surprise that it 
should not yet have attracted the attention of comparative psychologists. 
The series of actions to be performed once, and once only, in a lifetime bv 
an uninstructed, inexperienced atom is such that we should, a priori, have 
denounced it as an impossible means of existence, were it not shown that 
it is constantly successful. It is no wonder that the female Meloe produces 
five thousand times more eggs than are necessary to continue the species 
without diminution in the number of its individuals, for the first and most 
important act in the complex series of this life-history is accomplished by 
an extremely indiscriminating instinct; the newly hatched Meloe lias to 

Beetles 293 

get on to the body of the female of one species of bee; but it has no dis- 
crimination whatever of the kind of object it requires, and, as a matter of 
fact, passes with surprising rapidity on to any hairy object that touches it; 
hence an enormous majority of the young are wasted by getting on to all 
sorts of other insects; these larvae have been found in numbers on hairy 
Coleoptera, as well as on flies and bees of wrong kinds; the writer has ascer- 
tained by experiment that a camel's-hair brush is as eagerly seized, and 
passed on to, by the young Meloe as a living insect is." 

The commonest Eastern species of blister-beetles belong to the genus 
Epicauta. They feed when adult on the leaves of potato — being therefore 
often called potato-beetles — and on the pollen of goldenrod. E. pennsyl- 
vanica is uniformly black; E. cinerea is grayish black or even ashy, always 
with the margins of the elytra gray; E. vittata (Fig. 401) is yellowish or reddish 
above, with head and prothorax marked with black and with two black stripes 
on each elytron. In Meloe the wings are lacking and the elytra short and 
diverging; M. angusticollis , the buttercup oil-beetle, i to f inch long, 
of violaceous color, is the commonest eastern species. In the west the 
commonest blister-beetles are metallic green and blue and belong to the 
genus Cantharis. 

Another small family of rarely seen heteromerous beetles, which, how- 
ever, possess an extremely interesting and wonderfully specialized life-history 
and show a marked degenerate structure due to their parasitic habits, is 
the Stylopidae, or wasp parasites. Indeed these 
curiously modified beetles difl'er so much from 
all the other Coleoptera that some entomolo- 
gists look on them as composing a distinct order 
which these naturalists call Strepsiptera. The 
males are minute with large fan-shaped wings 
and reduced, short, club-like elytra. The 
females are wingless and never develop bevond 

a larval or grub-like condition. They live in Fig. 403.-A wasp Po//5/r5 sp., 
° -' parasitized by (x) Xeiws sp. 

the body of a wasp or bee (Fig. 403) — certain (After Jordan and Kellogg; 

foreign species parasitize ants, cockroaches, and slightly enlarged.) 
other insects — while the free-flying males live from only fifteen or twenty minutes 
to a day or two: three days is the longest observed lifetime of active adult 
existence! The youngest larva of the Stylopids — the egg-laying has not 
been observed — is a minute, active, six-legged creature, not unlike the Meloid 
triungulin, which attaches itself to the larva of a bee or wasp and burrows 
into its body. There it Hves parasitically, meanwhile undergoing hypermeta- 
morphosis in that after its first moult it becomes a footless maggot or grub. 
In this state it continues until, if a male, it pupates in the host's body and 
issues for its brief active adult Ufe. If a female, there is no pupation, but 

294 Beetles 

when the host larva itself pupates the Stylops pushes one end of its own 
body out between two abdominal segments of the host, and there gives birth 
alive to many little triunguHns. How the triungulins find their way to 
their bee-larva hosts is not very clear, but they probably he in wait in flowers 
and when a bee comes along they cHng to its leg and are thus carried to 
the nest where the larvae are. There are two genera of Stylopidse in our 
country, Xenos, which parasitizes the social wasps, Polistes, and Stylops, 
which parasitizes the mining-bees, Andrena. The triungulins of Xenos, 
being born in a community nest, can simply roam about over the brood- 
comb until they they find a wasp-larva to burrow into. 


In this suborder are included all those beetles known as curculios, wee- 
vils, bill-bugs, and snout-beetles (excepting the pea- and bean weevils, see 
p. 281). They are all characterized by the peculiar prolongation of the 
front of the head into a beak or snout, which may be long, slender and 
curved, or straight, short, thick, and obtuse. The mouth-parts, of which the 
small sharp jaws are the conspicuous feature, are situated at the tip of the 
snout; upper lip (labrum) and palpi are wanting. The antennae arise 
from the sides of the snout and are angularly bent or "elbowed" in the 
middle and end in a knobbed or clavate tip. The body is soHd and compact, 
usually strongly rounded above, and many species are thinly or thickly cov- 
ered with scales. 

Most of the weevils feed, as adults, on fruits, nuts, and various seeds, 
though some attack stems and leaves, and others hard wood. Many 
feign death when disturbed, folding up their legs and head and lying 
inert until danger is past. The larvae are soft, wrinkled, white, footless 
grubs which mostly live in fruits, nuts, and seeds. The larvae and adults 
of the important family Scolytidae, variously called timber-beetles, bark- 
borers, or engraver-beetles, burrow in the bark and wood of trees living or 

The principal families of the suborder can be separated by the following 

The dorsum of the last segment (pygidium) of the male divided transversely, so that 
this sex appears to have one more body-segment, when viewed dorsally, than 
the female. 
Mandibles with a scar on the anterior aspect. 

(Scarred snout-beetles.) Otiorhynchid.e. 

Mandibles without scar on the anterior aspect (Curculios.) Curculionid-i:. 

Pygidium of both sexes undivided. 
Pygidium vertical; tibis not serrate. 

(Bill-bugs and gianary-weevils.) Calandrid^. 
Pygidium horizontal ; tibiae usually serrate (B ark-beetles.) Scolytid^. 

Beetles 295 

The scarred snout-beetles, Otiorhynchidae, get their vernacular name 
from the presence of a distinct little scar on the front aspect of each mandible. 
It is made by the falling off of a mandibular appendage present in the pupa. 
Most of these beetles are covered with minute scales, much like those of the 
moths and butterflies, which give them often a bright metallic coloration. 
Several species of the family are injurious to fruits. 

The imbricated snout-beetle, Epiccerus imbricatiis, J inch long, dull 
silvery white with darker markings, and with the elytra with longitudinal 
lines of deep pits, has the posterior ends of the elytra very steep and cut off 
almost squarely and ending in a pointed process. It feeds on various culti- 
vated plants, as garden vegetables, strawberries, etc., and gnaws holes in 
the twigs and fruits of apple and cherry. The pitchy-legged weevil, 
Otiorhynchus ovatus, ^ inch long, dark brown to black with deeply pitted 
thorax and striated elytra, with deep punctures in the striae, almost egg- 
shaped hind body, and thorax with projecting angle on each side, attacks the 
roots and crowns of strawberry-plants, and also the leaves of apple-trees. 
Fuller's rose-beetle, Araniiges julleri, is perhaps the most familiar species 
of this family, as it attacks garden and conservatory roses, and in Cali- 
fornia is an orange pest of some note. It is \ inch long, oval, smoky-brown, 
and thinly covered with scales; its "snout" is short and obtuse. The eggs 
are laid in masses in concealed places on rose-bushes, 
the larvie feeding on the roots of the bushes, while the 
adults attack the leaves, buds, and flowers. The beetles 
hide during the day on the under side of the leaves, 
and can readily be collected and destroyed. 

The Curculionidae, the typical curculios and weevils, 
compose the largest and most important family of the 
suborder, comprising over 600 species of North Amer- 
ican beetles, and including many seriously destructive 

pests. Such enemies of the fruit-grower as the plum- ^ ^, 

^ ,. , , ^, , , Fig. 404.— The chest- 

curculio, plum-gouger, apple-weevil, and strawberry- nut-weevil, Balani- 

weevil, and such a destructive pest of cotton as the "'" caryatrypes. 

boll-weevil (for the study and combating of which natural sizeT' 

Congress has recently appropriated $250,000), are alone 

sufficient to give this family a high rank in the list of notorious insect 

pests. The eggs of Curculionids are laid singly in holes bored or cut by 

the female with her snout in stems or fruits of the food-plant and pushed 

to the bottom by the snout, which is therefore often very long and slender. 

The nut- and acorn-weevils of the genus Balaninus are characterized by 

their possession of an unusually long, slender, curving beak (Fig. 404) ; in 

the females this beak may be twice as long as the rest of the body; in the 

males it is usually about the length of the body. These beetles are from 

296 Beetles 

^ to I inch long, clay-yellow or mottled brownish, and lay their eggs in 
chestnuts, hazelnuts, acorns, walnuts, hickory-nuts, etc. The white, yellow- 
headed, maggot-like larva feeds on the kernel, and is full-grown at the 
time the nuts drop. It either lies in the nut over winter or crawls out and 
into the ground, where it pupates, and transforms into an adult; B. rectus 
and B. quercns are common acorn-weevils, B. caryatrypes (Fig. 404) a 
common chestnut-weevil, and B. nasicus a hickory-nut weevil. 

The genus Anthonomus includes small pear-shaped, modestly colored 
weevils with long slender snouts. A. qitadrigibbus, the apple-weevil, ^ inch 
long, dull brown, with four conspicuous brownish-red humps on the hinder 
part of the body, lays its eggs in little blackish-margined holes drilled into 
apples; the white, footless, wrinkled, brown-headed larva on hatching bur- 
rows into the core, feeds around it, ejecting much rusty-red excrement, and 
finally pupates, the adult weevil gnawing its way out to the surface. A . sig- 
natiis, the strawberry-weevil, blackish with gray pubescence, punctures 
the buds, laying an egg in each, and then punctures the flower-pedicel below 
the bud, so that it drops off; the larva feeds on the fallen unopened bud, 
changing to a beetle in midsummer. A. grandis is the notorious boll- 
weevil of the South, which has made its way since 1890 from Mexico into 
this country and is now one of our most serious insect pests; it destroys as 
much as ninety per cent of the cotton-crop in badly infested localities. The 
eggs are deposited in the buds and bolls, and the larva? feed on seed and 
shell, pupating inside the wall of the boll, through which the issuing beetle 
gnaws its way. This pest seems to feed only on cotton. 

Next to the codlin-moth and San Jose scale probably the most notorious 
and destructive frviit-pest is the plum-curculio, Conotrachelus nenuphar 

(Fig. 405), a small beetle, \ inch long, brown, 
and with four small elevated excrescences on 
the hard wing-covers. The beetles hibernate 
in rubbish, such as accumulated leaves, about 
the orchard, and come out in early spring to feed 
on the tender buds, leaves, flowers, and even 

^ ^, , ,. green bark. When the plums have set, the 

Fig. 40s. — The plum-curcuho, '^ . ... . , 

ConStrachelus nenuphar, females begm to deposit their eggs m them by 

(After photograph by Slinger- driUing a. tiny hole and pushing an egg into 

, en arge . each. Then a concentric slit is cut near the 

hole so as to leave the egg in a little flap in which the tissue is so injured 

that the rapid growing of the fruit does not injure the delicate egg buried 

in it. The whitish larva bores in until it reaches the stone around which 

it feeds. (The larva of the plum-gouger, Coccotorus sciitellaris , another 

destructive Curculionid pest of the plum, bores into the stone.) When 

the larvae are full-grown the infested plums fall to the ground, and the larvae 



crawl out and into the soil to pupate. The adult beetles soon issue and 
hunt up hibernating quarters. The plum-curculio attacks cherries, and 
also peaches, nectarines, and apricots. In many regions of this country 
it has wholly stopped the growing of plums. Curiously enough, but 
fortunately, this pest does not seem to be able to maintain itself in California, 
where plum (prune) growing is one of the chief industries. A remedy of 
some effectiveness is to jar each plum-tree, under which a sheet has been 
spread, repeatedly during blossoming and fruit-setting time. The curculios, 
alarmed by the jarring, fold up their legs and snout and fall to the ground 
(sheet), where they feign death. This feigning can be turned into reality 

Fig. 406. — Larva and pupa of the quince-curculio, Conotrachelus cratmgi. (After photo- 
graphs by Slingerland; at left, larva, natural size and enlarged; at right, pupa much 

by any one of various means. Excellent "curculio-catchers" consist of 
wheelbarrows on each of which is mounted a large inverted umbrella split 
in front to receive the tree-trunk, against which the barrow (with a padded 
bumper) is driven with force enough to do the jarring. All fallen plums also 
should be promptly gathered and burned or scalded so as to kill the larvae 

The family Calandridas includes about eighty North American species 
of weevils, of which several are common and famihar under the names of 
corn bill-bugs and rice- and grain-weevils. To the large genus Sphenophorus 
belong the species known as corn bill-bugs, blackish, brown, or rarely gray 
in color, from i to J inch long, with thick and hard elytra which are 
ridged and punctured, as is also the thorax. By day they hide in the soil 



at the base of young corn-plants, and at night bore little round holes into 
their stems. The larvae live in the stems of timothy, sedges, or bulb-rooted 
grasses, pupating in fall or early spring. To the genus Calandra belongs 
the destructive rice-w^eevil, C. oryza, \ inch long, blackish to pale chestnut, 
which attacks all kinds of stored grains and is especially injurious in the 
southern states to rice, and the granary- weevil, C. granaria, ^ inch long, 
dark brown, also common in grain-bins. Both these species have been 
widely distributed by commerce, and by their rapid multiphcation and the 
concealment afforded them by the grain often attain such abundance as 

to cause great loss in mills, breweries, 
and elevators. The preventive remedy 
is cleanliness and the rapid removal of 
the stored grain. They prefer dark 
places, therefore a flood of sunHght 
will prevent their rapid increase. In 
bins that can be made nearly air-tight 
these pests may be killed by the fumes 
of carbon bisulphide. 

One may often see in the woods the 
curious hieroglyphics of the engraver- 
beetles (Scolytidae). Where bark has 
been torn from a tree - trunk both 
the exposed trunk-wood and the inner 
surface of the stripped-off bark reveal 
the tortuous branching mines or tunnels 
of the Scolytidae. A common way of 

„ . 1- /- making these tunnels is as follows: The 

Fig. 407. — ^The quince -curculio, Cono- & 

trachelus cratagi. (After photograph beetles (a male and a female together) 
by Slingerland; natural size and en- burrow from the outside through the 
larged.) , . , , , , „ T • 

thick rough outer bark, usually leaving 

a little betraying splotch of fine sawdust, to the inner live bark or sap- 
wood; here the pair turn, keep to this Uve sap-filled region, laying their 
eggs in masses or scattered along a tunnel. Soon the larvae hatch, where- 
upon each digs a tunnel for itself, all of the new larval mines branching out 
from the original tunnel made by the parent beetles. When full-grown 
the larva digs a cell at the end of its tunnel and pupates in it. The issuing 
beetle finds its way out through the tunnels and is soon ready to begin a new 
mine. But there is much variation in the mining habits of the various species. 
The beetles are small, often microscopic, the larger ones rarely more than 
\ inch long. They are brown to blackish, with stout, nearly cylindrical 
hard bodies, the hind end of the body usually obliquely or squarely truncate, 
and the head short, bent downward; and so covered by the thorax as to be 



almost invisible from above. The larvK are white and footless little grubs 
with very strong jaws. The family includes 1 50 species in North America, and 
because of the recently awakened interest in forestry is now being given special 
attention by entomologists. The losses, by the death of trees and the rid- 
dling of timber, caused by these obscure little insects are enormous. Pinchot, 
chief of the United States Bureau of J'orestry, has recently estimated the 
annual forest losses caused by insects to be $100,000,000, and most of the 
ravages are due to the Scolytidae. 

Among the most destructive genera are Dendroctonus and Tomicus, 
each with numerous species. They often work in the same tree. For 
example, the famous Monterey pines of California are attacked by Dendroc- 

FiG. 408. — Galleries in Monterey pine, with larvas, pupae, and adults of the engraver- 
beetle, Tomicus plaslographus. (Natural size except the single beetle outside, 
which is enlarged three times.) 

tonus valens in the lower three or four feet of the trunk, as many as four 
hundred individuals (larvae, pupae, and adults) occurring in this limited 
space in badly infested trees, while above this zone on up to the top of the 
tree are the mines of Tomicus plaslographus (Fig. 408), from thirty to forty 
pairs burrowing into each yard of trunk. It is plain that such a combined 
attack on a single tree means death to it. 

The ambrosia-beetles, including half a dozen genera and many species, 



have special habits which make them comparable in some ways with the 
social wasps, bees, and ants, and with the termites. They live in mines — 
the "black holes" often seen in timber — bored into the heart-wood of sick 
or dead trees, in colonies including numerous adults and many larvae. Their 
food is not the wood of the tree, but consists of certain minute and succulent 
bodies produced by a fungus which grows on the walls of their burrows. 
This fungus does not grow there by chance, but is "planted" by the beetles. 
It is started by the female upon a carefully packed bed or layer of chips, 
sometimes near the entrance of a burrow, in the bark, but generally at the 
end of a branch gallery in the wood. It spreads, or is spread, from this 
forcing-bed to the walls of the various galleries and chambers of the mine. 
The young larvae nip off the tender tips of the fungus stalks "as calves crop the 
heads of clover," but the older larvae and adult beetles eat the whole structure 
down to its base, from which new hyphae soon spring up afresh. The fungus 
is suitable for the insects only when fresh and juicy: if allowed to ripen, the 
tender protoplasm is shut up in spores, and the galleries are soon filled to 
suffocation with these spores and the ramifying mycelial threads. Indeed 
the colony of ambrosia-beetles — ambrosia being the name applied to the 
tender fungus food — is often overwhelmed and destroyed by the quick 
growth of their garden-patch. If anything happens to interrupt the constant 
feeding on and cutting back of the fungus, the colony is almost always 


EXT to the name ''bug" there is no other name so 
popular in point of miscellaneous application to insects 
as "fly." This looseness of popular nomenclature 
may be largely due to the fact that entomologists them- 
selves apply the term "fly " in several compound words, 
as butterfly, alder-fly, caddis-fly, May-fly, saw-fly, and 
the Hke, to widely differing kinds of insects. Used as 
a simple word, however, by fly an entomologist means 
some species of the order Diptera. The various kinds of 
true flies have of course special names, as mosquitoes, 
midges, punkies, gnats, or as in the compounds 
horse-flies, bee-flies, flower-flies, robber-flies, etc. 
The order Diptera is so large and includes insects of such widely differing 
form and habit that it is difficult to formulate any general account of it. The 
gj—ju . name itself is derived from the most conspicuous 

structural condition of flies, namely, their two- 
winged state. All Diptera have but a single 
pair of wings, if any; a few are wingless. The 

tiG. 409. Fig. 410. 

Fig. 409. — Mouth-parts of a female mosquito, Culex sp. lep., labrum-epipharynx; md., 
mandible; mx.L, maxillary lobe; ntx.p., maxillary palpus; hyp., hypopharynx; li., 
labium; gl., glossa; pg., paraglossa. 

Fig. 410. — Mouth-parts of the house-fly, Miisca domestica. lb., labrum; mx.p., maxil- 
lary palpi; li., labium; la., labellum. 

hind wings of other forms are replaced by a pair of strange little structures 



The Two-winged Flies 

Fig. 411. — Head, antennae, 
and beak of mosquito, lat- 
eral aspect. 

called balancers, or halteres, whose use seems to be chiefly that of orienting 
or directing the fly in its flight. The possession of these balancers 
is a certain diagno.stic character in distinguishing Diptera from all 
other insects. The wings are membranous and usually clear, and 
supported by a few strong veins. No flies can bite in the sense 
of the chewing or crushing biting common to beetles, grasshoppers, and 
other insects with jaw-like mandibles, but some have mandibles elongate, 
slender, and sharp-pointed, so that they act as needles or stylets to make 
punctures in the flesh of animals or tissues of plants. The great majority 
of flies, however, have no mandibles at all and no piercing beak, but lap up 

liquid food with a curious folding fleshy proboscis, 
which is the highly modified labium or under-lip. 
They feed on flower-nectar, or any exposed sweet- 
ish Hquid, or the juices of decaying animal or 
plant substance. To take solid food as the 
house-fly does from a lump of sugar, the solid 
has to be rasped off as small particles which are 
either dissolved or mixed in a salivary fluid 
that issues from the fleshy tip of the proboscis. 

All the Diptera have a complete metamorphosis, the young hatching 
from the egg as footless and often headless larvae (maggots, grubs), usually 
soft and white, and in many cases ob- 
taining food osmotically through the 
skin. The life-history is usually rapid, 
so that generation after generation suc- 
ceed one another quickly. Thus it may 
be true, as an old proverb says, that 
a single pair of flesh-flies (and their 
progeny) will consume the carcass of 
an ox more rapidly than a lion. The 
pupae of the more specialized flies are 
concealed in the thickened and darkened 
last larval moult, the whole puparium 
looking much like a large elliptical brown 

The Diptera include the famihar 
house-flies, flesh-flies, and bluebottles 

of the dwelling and stables; the horse-flies and greenheads, that make 
summer hfe sometimes a burden for horses and their drivers; the buzzing 
flower- and bee-flies of the gardens; the beautiful little pomace-flies with 
their brilliant colors and mottled wings that swarm like midges about 
the cider-press and faUen and fermenting fruit; the bot-flies, those disgust- 

FiG. 412. — The blow-fly, Cclliphoraery- 
throcephala. Larva, pupa, and adult. 

The Two-winged Flies 303 

ing and injurious pests of horses, cattle, rabbits, rats, etc. ; the fierce robber- 
flies that prey on other insects, including their own fly cousins; the midges 
and gnats, that gather in dancing swarms over pastures and streams; the 
black-flies and punkies, dreaded enemies of the trout-fisher and camper; 
and, worst of all, the cosmopolitian mosquito, probably the most serious insect 
enemy of mankind. Only in recent years have we come to recognize the 
mosquito's real capacity for mischief. Annoying and vexatious they have 
always everywhere been, by day and night, from tropics to pole, from the 
salt marshes by the sea to the alpine lakes on the shoulders of the mountain- 
peaks. But that the mosquito-bite not only annoys but may kill, by infect- 
ing the punctured tissues with the germs of malaria or yellow fever or filari- 
asis, three of the most wide-spread and fatal diseases of man — this alarming 
fact is a matter which has come to be really recognized only recently, and 
the general recognition of which has given to the practical study of insects 
an importance which years of warning and protesting by economic entomol- 
ogists have been wholly unable to do. 

The Diptera include about 7,000 known species in North America, thus 
ranking among the principal orders of insects in degree of numerical represen- 
tation in this country. About 50,000 species are known in the whole world. 

The order may be separated into certain principal subdivisions by the 
following table: 

Living as external parasites on mammals, birds, or honey-bees; body flattened and 
often wingless; the young born alive as larvae nearly ready to pupate. 

Suborder Pupipara (see p. 351). 
Not living on the bodies of other animals; young usually produced as eggs. 

Suborder Diptera genuina (see p. 304). 

Antennae with numerous (more than five) segments. .Section Nematocera (see p. 304). 

Antennae with not more than five segments, usually with three, the third sometimes annu- 

lated, showing it to be a compound segment, i.e., composed of several coalesced 

segments Section Brachycera (see p. 327). 

Third segment of antennae annulated, showing it to be composed of several coalesced 

segments (see p. 327). 

Antennae consisting of four or five distinct segments (see p. 330). 

Antennae with but three segments (rarely less), the third segment with or without a 
style or bristle (see p. 332). 

Of the two suborders the smaller one, the Pupipara, including certain 
strangely speciahzed and degraded parasitic flies, will be considered last. Of 
he first suborder, the Diptera genuina, the various famihes of small midge- 
and mosquito-Hke flies composing the section Nematocera (flies with slender 
several-segmented antennas) will be discussed first, as they are believed by 
entomologists to be the more generalized or simpler flies. 

304 The Two-winged Flies 

Of this section the mosquitoes, black flies, and punkies are perhaps best 
known because of the annoyance and irritation caused by their "bites," 
that is, the punctures made by the sharp beak of the females in their blood- 
sucking forays. But the swarms of dancing midges and the sprawhng long- 
legged crane-flies, or leather-jackets, are not unfamiliar members of this group. 
In addition there belong here a few famiHes of flies httle known but possessed 
of most interesting habits and form. 


(The references to the names and character of the veins in the wings which occur in 
this and other keys used in this chapter may be understood by a comparison of the 
venation of the specimen being examined with Fig. 18, and with the figures of the 
venation of various families, as Figs. 425, 436, 444, etc.) 

A. Antennae slender, longer than thorax; usually nearly as long as body or longer; 
legs long and slender, and abdomen usually so. 

B. Very small moth-like flies, with body and wings hairy; wings with 9-1 1 longi- 
tudinal veins, but no cross-veins except sometimes near the base of the wing. 

(Moth-like flies.) Psychodid^. 
BB. Not as above. 

C. Wings with a network of line vein-like lines near the outer and hinder 

margins in addition to the regular (heavier) venation. 

(Net-winged midges.) Blepharocerid^. 
CC. The margin of the wings and the veins fringed with scales. 

(Mosquitoes.) Culicid^. 
CCC. With a distinct V-shaped suture on the back of the thorax. 

(Crane-flies.) Tipulid^, 
CCCC. Without distinct V-shaped suture on the back of the thorax. 

D. Anal veins entirely wanting; medial vein wanting or at most 
represented by a single unbranched fold. 

(Gall-gnats.) Cecidomyiid^. 

DD. Anal veins present or represented by folds; medial vein present 

or at least represented by a fold which is usually branched. 

E. Ocelli present; hgs slender and with greatly elongate 

ccxse (basal segment). . . (Fungus-gnats.) Mycetophilid^. 

EE. Ocelli absent. 

F. Wing-veins well developed in all parts of the wing. 

(Dixa-flies.) DixiD^. 
FF. Wing-veins much stouter near the costal (front) 
margin of the wing than elsewhere. 

(Midges.) Chironomid^. 
AA. Antennse shorter than the thorax and rather stout. 

B. OcelH present (March-flies.) Bibionid^. 

BB. Ocelli absent; wings very broad (Buffalo-gnats.) Simuliid^. 

Of the ten families included in the above key the members of five pass 
the young stages, larval and pupal, in fresh water; of the members of two 

The Two-winged Flies 


some have aquatic immature stages and some terrestrial; while the larvae 
and pupaj of all the members of the remaining three live in plants or in the 
ground, none being aquatic. 

Best known of the aquatic famihes, and indeed of the whole suborder, 
is the mosquito family, the 
Culicidc'e. While the different 
kinds of mosquitoes are much 
alike, so much so indeed that 
most of us are quite content if 
we can determine an insect to be 
a mosquito without carrying the 
identification farther, there are 
known in the world at least 300 
different mosquito species, rep- 
resenting two dozen distinct 
genera. In North America 
nearly 60 species are already 
known, representing 10 genera, 
and new ones are being found 
constantly. In the family Culi- 
cida; are included two distinct 
general types of mosquito, one 
with mouth-parts forming a long, 
slender, sucking proboscis, pro- 
vided with sharp, needle-like 
stylets for piercing (Fig. 411), the 
other with the mouth-parts short 
and better adapted for lapping 
or sucking up freely exposed 
liquids. The latter type of 
mouth is possessed by but two 
genera, all the others being 
piercers and blood suckers (in 
the female sex). Of these pierc- 
ing genera three are of especial 
importance and interest to us 
because of their abundance and 

their definitely determined relation to the development, incubation, and dis- 
semination of certain serious diseases of man. These three genera are Culex, 
Stegomyia, and Anopheles. To Culex belong the great majority of familiar 
mosquitoes which pursue and harass us with their songs and bites; to Stego- 
myia (and Culex) belong the mosquitoes held responsible for the dissemination 

Fig. 413. — The life-history of a mosquito, 
Culex sp A small raft of eggs is shown on 
the surface of the water, several larvae 
("wrigglers"), long and slender, and one 
pupa ("tumbler"), large-headed, are shown 
in the water, and an adult in the air above. 
(From life; much enlarged.) 

306 The Two-winged Flies 

of yellow fever and filariasis, and to Anopheles belong the malaria breeding 
and distributing mosquitoes. 

All the mosquitoes agree in having strictly aquatic immature stages. The 
eggs are laid on the surface of standing or slowly moving water, usually 
fresh, although several species breed abundantly and probably exclusively 
in brackish water. These eggs are in small one-layered packets or rafts (usual 
in Culex) (Fig. 413) or are scattered singly (in Stegomyia and Anopheles) (Fig. 
414) and hatch in from one to four days, varying with the species, and in 
the same species with the temperature and hght 
conditions. The water oviposited on may be, for 
Culex, that of a pond, a pool, or any temporary 
puddle, or even that in an exposed trough, barrel, 
pail, or can. With Anopheles only natural, usually 
Fig. 414.— The eggs of permanent, pools are selected. I have found the 
Anopheles sp. (After gggg of Culex incidens on the surface of a bubbhng 
Giles; much enlarged.) g^^^^.^pj-jng jn CaHfornia, and of Stegomyia in water 
held in sUght depressions in a number of ship's metal parts in Samoa. 
The brackish-water species of Culex usually lay their eggs on the small 
clear pools scattered through the marshes. A few entomologists have 
recorded their belief, based on various indirect observations, that the eggs 
of Anopheles at least may be deposited on the soil, but no direct proof of 
this is yet on record. 

The larvs (Figs. 413 and 415) of mosquitoes are the familiar wrigglers of 
ponds and ditches. The long, slender, squirming body, with its forked posterior 
extremity and thick head end, is thoroughly characteristic. The head is 
provided with a pair of vibratile tufts or brushes of fine hairs which are 
kept, most of the time, in rapid motion, creating currents of water setting 
toward the mouth, and thus bringing to it a constant supply of food, which 
consists of organic particles and microscopic animals. Breathing is accom- 
plished by the wrigglers coming to the surface and hanging head downward 
from it with the open tip of the respiratory tube, one of the prongs of the 
posterior forking of the body, projecting just through the surface film. If a 
mosquito wriggler is prevented from coming to the surface, or if, once there, it 
finds some impediment which restrains it from getting its respiratory tube 
into connection with the free air above the surface, it will drown. And 
this fact partly explains the fatal effectiveness of a film of kerosene spread 
over the surface of a pool in which mosquitoes are breeding. The larval 
stage lasts from one to four weeks, varying in different species and also 
varying in the case of each species at different seasons and under different 
conditions of food-supply, temperature, and light. Larva? of Culex have 
lived in breeding-jars in my laboratory for three months. The larvae moult 
twice, and on the third casting of the skin appear as active, non-feeding 

The Two-winged Flies 307 

pupae (Figs. 413 and 415) with thick, broad head end (the thick part includes 
thorax and head) and slender, curving abdomen, bearing two conspicuous 
swimming-iiaps at the tip. The pupa rests at the surface of the water with 
its two short horn-like respiratory tubes, which rise from the dorsum of the 
thorax, extending through the surface film to the air above. When dis- 
turbed it swims swiftly down into the water by quick bendings or flappings 
of the abdomen with its terminal flaps. The pupal stage lasts from two to 
five days, with comparatively little variation beyond these extremes. 

The adults issue through a longitudinal rent in the back of the pupal 
cuticle, and while drying their wings, legs, and body vestiture rest on the 
surface of the water, often partly supported by the floating discarded skin. 
The two wings are long and narrow, the legs long and slender, the thorax 
humped with the small head hanging down in front and the slender sub- 
cylindrical abdomen depending behind. The body is clothed with scales, as 
are the veins of the wings, and on the scales, which are of different shapes 
and sizes on different parts of the body, and vary in different species, depend 
the colors and pattern, often striking and beautiful, just as all the color pat- 
terns of the butterflies and moths are produced by a covering over body and 
wings of similar scales. The males of all mosquitoes differ from the females 
in having the slender, many-segmented antennas provided with many long 
fine hairs arranged in whorls and combining to give the antennae a bushy or 
feathery appearance. . These hairs, as has been proved by experiment and 
histologic study, are a part of an elaborate auditory apparatus, their special 
function being to be set into vibration when impinged on by sound-waves of 
certain rates of vibration, and to transmit this vibration to a complex nervous 
organ in the second antennal segment (Figs. 56 and 57). The males, while 
having a long, slender, sucking-proboscis, do not possess the piercing sty- 
lets characteristic of the female, and hence are not blood-suckers, but prob- 
ably feed, if at all, on the nectar of plants or on other exposed liquids. The 
females suck blood when they can get it, but in Heu of this animal fluid 
feed on the sap of plants. In experimental work in the laboratory cut 
pieces of banana are provided the imprisoned adult mosquitoes. 

At this writing about fifty species of Culex, one species of Stegomyia, and 
four species of Anopheles have been found in this country. These three 
genera may be distinguished by the following key: 

Palpi (the mouth-feelers projecting by the side of the proboscis) long in both male and 

female, about as long as the proboscis Anopheles. 

Palpi as long as proboscis in male, but only one-third as long in female. 

Scales on the head narrow and curved Culex. 

Scales on the head flat and broad Stegomyia. 

Our particular interest in being able to distinguish these genera lies, as 
already said, in the special relation which their members bear to certain 


The Two-winged Flies 

wide-spread and serious human diseases. The role played by mosquitoes 
in the breeding and dissemination of the microscopic germs of malaria has 
been so well exploited in newspapers and magazines that, although a matter 
of comparatively recent determination, it is already common knowledge, at 
least in its more general outline. For a somewhat detailed account of the 
etiology of the diseases known to be disseminated by mosquitoes, including 
the exact relation of the mosquito host to the disease-germs, see Chapter 
XVIII of this book. It is sufficient to say here that the malarial germs seem 
to live parasitically in and be disseminated by the various species of Ano- 
pheles only, the yellow- fever germs only by the species Stegomyia jasciata, and 
the minute worms of filariasis by the same species and two or three tropical 
forms of Culex, while the score and more of North American species of 

Fig. 415. — A malaria-carrying mosquito, Anopheles maculipennis ; larva at left, in 
middle two eggs below and pupa above, male adult at right. (From life; much 

Culex compose most of the hordes of piercing and blood-sucking mosqui- 
toes which in so many localities make life distressful. Stegomyia jasciata is 
found in this country only in the Gulf states. In our colonies, the Hawaiian 
and American Samoan Islands, I have found it to be the most abundant mos- 
quito species, although yellow fever is yet unknown in these islands. But 
it seems not improbable that, with the cutting of a canal through the Isthmus 
of Panama so that ships can sail directly from the West Indies to Hawaii 
continuously within the tropics, Stegomyia individuals infested with yellow- 
fever germs might be readily carried to our tropical Pacific colonies. Such 
a possible contingency should at least be had in mind by those charged with 
the responsibility of public-health affairs in Hawaii and Samoa. Stegomyia 
is already terrible enough in its disease-spreading capacity in unfortunate 
Samoa, as explained in Chapter XVIII, the frightful scourge elephantiasis, 

The Two-winged Flies 


an incurable and hideously deforming kind of filariasis, from which quite 
one-third of the natives of Samoa suffer, being disseminated chiefly (so 
far as our present knowledge permits us to affirm) by mosquitoes of the 
species Stegomyia fascia ta. 

With a few English investigators and our own government and state 
entomologists in the lead, a great campaign is being waged against mos- 
quitoes. Despite the hosts of the enemy, its great capacity for providing 
new individuals to supply the places of the fallen, i{s effective means of 
locomotion, and its easily managed de- 
partment of commissary, local foraging 
being exclusively relied on for sustain- 
ing its armies, we are making headway 
against it. Our modes of attack are 
various: by draining swamps, ponds, 

Fig. 416. Fig. 417. 

Fig. 416.— a short-beaked mosquito, Corethra sp. (From life; four times natural size. 
Fig. 417. — Pupa (at left) and larva (at right) of short-beaked mosquito, Corethra sp. 
(From life; six times natural size.) 

and puddles we restrict the multipHcation of these pests, and rid particular 
localities of them altogether; by introducing into ponds and pools which 
cannot be drained substances, as kerosene, etc., which are poisonous to mos- 
quitoes, we kill them in their adolescence; by encouraging and disseminating 
their natural enemies, such as dragon-flies, we pursue them in their own 
elements, water and air. Mosquitoes do not fly far; when abundant in a 
locality, breeding-places are to be looked for close at hand. The open rain- 
water barrel, a little puddle by the lawn hydrant, a cistern with unscreened 
openings, all of these are welcome invitations to the mosquito to come and 
rear a large family. Put close screen tops over water in cisterns and barrels; 



The Two-winged Flies 

leave no standing puddles in the back yard or decorative lily-pools in the 
front; pour kerosene on the surface of ponds and ditches in the neighbor- 
hood, and the mosquito problem for localities not adjacent to swamps and 
marshes is nearly solved. Where the problem includes swamps larger 
measures must be undertaken, community effort may be necessary, and the 
rhunicipal or county administration called on to take official action. But 
when it is remembered that aboHshing the mosquito pest means doing away 
with malaria, and in the subtropic and tropic region with yellow fever and 
filariasis, no pains will seem too troublesome, no expense too large in this 
warfare of man against mosquitoes. 

Fig. 418. Fig. 410. 

Fig. 418. — Scales on the wings of Culex fatigans. (After Theobald; greatly magnified.) 
Fig. 419. — A midge, male, Chironomiis sp. (From life; much enlarged.) 

Looking not unlike mosquitoes are the larger species of the family Chiro- 
nomidae, whose members are popularly known as midges and punkies, the 
name blood-worm being applied to the reddish aquatic larvae of certain 
species. Like the mosquitoes, the males are distinguished from the females by 
their very bushy or feathery antennae, but, unlike the mosquitoes, the females, 
except in the case of the minute punkies or "no-see-ums" of the New Eng- 
land and Canadian mountains and forests, and their near relatives in the 
western forests, are not blood-suckers. The midges are particularly notice- 
able in "dancing-time," that is, when they collect in great swarms and toss up 
and down in the air over meadows, pastures, and stream sides. 

The larvae (Fig. 420) of most species are aquatic, some of them forming 
small tubular cases, as caddis-fly larvae do, and most oi them being distinctly 
reddish in color. They wriggle about in the slime and decaying leaves at 
the bottom of ponds or lakes, feeding on vegetable matter. The pupae 
(Fig. 421) are, like those of the mosquitoes, active, although of course non- 
feeding, and are provided with two bunches of fine hair-like tracheal gills 
on the dorsum of the thorax, or with a pair of short club-shaped processes 

The Two-winged Flies 


which have a sort of sieve-like skin. In both cases the pupa breathes the 

oxygen which is mixed with water and is thus not 

compelled, as are the mosquito pupae, to come to the 

surface for air. The larvae of the genus Ceratopogon 

and its alhes, which include the fiercely biting and 

blood-sucking httle punkies (Fig. 422), so irritating 

to the fisherman and hunter in the north woods, 

Fig. 420. Fig. 421. 

Fig. 420. — Larva of a midge, Chironomus sp. (From life: natural length t inch.) 
Fig. 421. — Pupa of midge, Chironomus sp. (From life; natural length \ inch.) 

live, according to Comstock, "under the bark of decaying branches, under 
fallen leaves, and in sap flowing from wounded trees. " 

Running and half flying about over the spray- wet rocks and on the surface 
of the smaller tide-pools between tide-lines on the ocean shore near Mon- 

FiG. 422. Fig. 423. 

Fig. 422. — Mouth-parts of a female "punkie," Ceratopogon sp. lb., labrum; md., 

mandible; «;x., maxilla; ?«:x:./., maxillary lobe; m^x;./)., maxillary palpus; //., labium; 

p.g., paraglossa; hyp., hypothorax. 
Fig. 423. — The tide-rock fly, Eretmoptera browni. (Natural length ^ inch.) 

terey, California, may be seen in the winter months many small, long-legged, 
spider-Hke flies (Fig. 423) whose wings are reduced to mere oar-like veinless 
rudiments. The larvae and pups live submerged in the salt water of the 
outer and most exposed tide-pools, where the ocean water is held in shallow 
depressions in the rocks, and is changed many times daily by the dashing 
of the waves. Where the flies go when the tide is in and these rocks are 


The Two-winged Flies 

either whouy submerged or at least constantly dashed over by the breaking 
waves, I have not been able to determine; but the larvae and pupje cling 

Fig. 424. 

Fig. 424.- 
Fig. 425.- 

FiG. 425. 
(Four times natural size.) 

-A black-fly, Simuliiim sp. 

-Diagram of wing of black-fly, Simnliitm, showing venation. 

tight and secure in their rock basins to small but strong silken nets spun 
by the larvae. They rest on the under side of these nets, indeed are aknost 
enclosed in them as in a cocoon. This little fly is a most interesting insect 
because of its ocean-water habitat — very few insects live in salt water, and 


almost no others have so truly an ocean home, except 
the curious salt-water striders, Halobates (see p. 
197), which live on the surface of the ocean far out 
at sea. It is interesting, too, because of its structu- 
ral modifications, the atrophied wings, rudimentary 
balancers, etc., which set it off widely from all 
other flies. Its tide-pool habitat is undoubtedly the 
result of a slow migration and adaptation in the 
course of many generations on the part of some 
shore-inhabiting fly. There are many small flies 
which frequent ocean beaches and rocks, feeding on 

Fig. 426. — Larvae and pupas of Simidium sp. on edge of stream, May-fly on projecting 

twig. (After Felt.) 

The Two-winged Flies 


decaying seaweed, etc., and from among these this species has no doubt 
gradually worked its way out to the very verge of the shore-line, becoming 
gradually adapted in habit and structure to the conditions of its new 

^ Besides the mosquitoes and punkies a third kind of fly assails the rod- 
and-Hne fisherman, the hunter, and the camper in forests and along the streams; 
black, stout-bodied, hump-backed, short-legged, broad-winged flies (Fig. 
424) from one-sixth to one-fourth of an inch long, with short but strong 
piercing proboscis. These are black-flies, buffalo-gnats or turkey-gnats, as 
they are variously called, composing the small family Simuliidas, distributed 
all over this country, but especially abundant in the southern states, where 
they attack cattle so fiercely and in such great swarms that the animals are 
driven frantic and sometimes even killed by a violent fever produced by the 
terrible biting. 

The larvae (Fig. 426) are odd, squirming, slippery, little black "worms," 
which, clinging by the hind tip of the body, occur in dense colonies or patches 
on the smooth rock bed in shallow places 
of swift streams. The lip of a fall is a 

favorite place for them. The swift- f^ }^[fp. \ \md. 

running water constantly affords them 
an abundant air and food supply. The 
free or head end of the body is provided j^"" ^"^^ // 


Fig. 427- Fig. 428. 

Fig. 427.— Mouth-parts of female black-fly, Simuliiim sp. lep., labrum; hyp., hypo- 
pharynx; md., mandible; mx., maxilla; 7nxp., maxillary palpus; li., labium; pg., 
paraglossa. (Much enlarged.) 

Fig. 428.— Mouth-parts of larva of black-fly, Simulium sp. lb., labrum; ep., epipharvnx; 
md., mandible; mx., maxilla; mxp., maxillary palpus; mxl., maxillary lobe;' li., 
labium; hyp., hypopharynx. (Much enlarged.) 

with a conspicuous pair of freely movable brushes which collect food from 
the water. The cHnging to the rock is effected by means of silk spun 
from the mouth, and by the skilful use of silken threads the larva? can 
move about over the submerged rock bed without being washed away by 
the swift water. When ready to pupate, which is after about a month of 

3 1 4 The Two-winged Flies 

larval life (under favorable conditions of temperature and food-supply), the 
larva spins a little silken cornucopia-like cocoon (Fig. 426) fastened to the 
rock by the httle end, and often fastened by the sides to adjacent cocoons. 
The large free end is left open. In this cocoon it pupates, and after about 
three weeks the winged fly issues. The eggs are laid in patches on the rocks 

Fig. 429. — Longitudinal section of head of old larva of black-fly, Simulium sp., showing 
adult mouth-parts developing inside of or corresponding with the larval mouth- 
parts. /.OT(/., larval mandible; /.wix., larval maxilla; /./?., larval labium; /.c, larval 
cuticle; /.a., larval antenna ; i.w J., adult mandible; i.wx., adult maxilla; i./i., adult 
labium; i.d., adult hypoderm (cell-layer of skin); i.a., adult antennae; i.e., adult 
eye. (Much enlarged.) 

just below the surface of the water, or on the spray-dashed sides of boulders 
in the stream or on its margin. 

' In the same places where the Simulium larvae live, that is, on the smooth 
rock faces of stream bed and lip of fall under the thin apron of swift silver 
water of mountain streams, live also the curious flattened larvae (Fig. 430) of 
the net-winged midges or Blepharoceridae. This small family of interesting 
flies, comprising only eighteen species in the whole world, of which seven 
belong to this country, is one with which the general collector will hardly 
become acquainted unless he takes particular pains to do so. But the pains 
are well worth while, for they are not pains at all, but pleasures. In the first 
place, the larvce — and they must be looked for first, the winged flies being very 
rare, very retiring, and hardly distinguishable, until captured, from a number 
of other common and less interesting kinds — live only in the most attractive 
parts of the most attractive mountain brooks. I have found them in a tiny 
swift stream near Quebec, in two or three hillside brooks near Ithaca, 
N. Y., in roaring mountain torrents in the Rocky Mountains, and in similar 
plunging streams in the Sierra Nevada and Coast Range. Clinging by a 
ventral series of six suckers to the smooth shining rock bed, the short broad 

The Two-winged Flies 


larvae squirm slowly around, feeding on diatoms and other microscopic water 

\ A / . 

Fig. 430. Fig. 431. 

Fig. 430. — Larva of net-winged midge, Bihiocephala comstocki. At left, dorsal view; 
at right, ventral view, ant., antennae; l.p., lateral processes; t.g., tracheal gills; 
s., sucker. (Natural length, f to \ inch.) 

Fig. 431. — Cross-section of body of larva of net-winged midge, showing anatomical 
details of sucker and other parts, h., heart; al.c, alimentary canal; l.p., lateral 
process; v.c, ventral nerve-cord; r., rim of sucker; s., stopper of sucker; m.s.c, 
muscles for retracting sucker and contracting body; t., tendon at end of muscles. 
(Much enlarged.) 

organisms, and never suffering themselves to get into 
planted from the highly aerated swift water of the 
stream's center to the slow water of eddies or pools 
along the bank, they die very soon. When ready 
to pupate they gather in small patches, still keeping 
in the swift water, and each changes into a curious 
flattened, turtle-shaped, motionless, non-feeding pupa 
(Fig. 432) which is safely glued to the rock face by 
its under surface. The dorsal wall is thick and black, 
and projecting from it at the broad front head end 
is a pair of breathing-organs, each composed of three 
or four thin plate-like gills. When the fly is ready 
to emerge the pupal skin splits longitudinally along the 
back, and the delicate body pushes up through this 
slit, and through the shallow swift water until the 
wings can be outspread. All this is quickly done, 
the fly being enchained by its long legs, which cling 
to the pupal shell until it can fly away. But the 

slow water. Trans- 

FiG. 432. — Pupa, dorsal 
aspect, of net-winged 
midge, Bihiocephala 
comstocki. Note re- 
spiratory leaves on 
dorsum of prothorax. 
(Natural length, J inch.) 


The Two-winged Flies 

Fig. 433. Fig. 434. 

Fig. 433. — Net-winged midge, Bibiocephala elegantuliis, female. (Natural length of 
body, f inch.) 

Fig. 434. — Mouth-parts of female net-winged midge, Bibiocephala doanei. l.ep., labrum- 
epipharynx; ind., mandible; mx., maxilla; mxl., maxillary lobe; mxp., maxillary 
palpus; li., labium; pg., paraglossa; hyp., hypopharynx. (Much enlarged.) 

Fig. 435. — Heads of female (at left) and of male (at right) of net-winged midge, Bibio- 
cephala comstocki, showing division of eyes into two parts, the upper part with fewer 
and larger facets than the lower part. (Much enlarged.) 

The Two-winged Flies 


swift water works great havoc among the weak, soft-bodied emerging creatures. 
I have watched many flies issuing, and a large proportion of them get swept 
away and presumably drowned before they can get their wings unfolded 
and themselves clear of the torrent. It is an extraordinary life-history that 

Fig. 436. — Primarv venation of wing of net-winged midge, Bibiocephala comstocki. 
R^, R^, etc., branches of the radial vein. (Much enlarged.) 

these files have, and the great danger attending the transformation to the 
adult stage probably partly explains why the species are so few. It is an 
unsuccessful type of insect hfe; the family is probably becoming extinguished. 
Because the few living species are so widely distributed over the world — 


Fig. 437. Fig. 438. 

Fig. 437. — Diagram of cross-section of head through compound eyes of net-winged 
midge, Blepharocera capitata, female, o, ocelli; br., brain; o.l., optic lobes; /./., large 
facets; s.}., small facets. 

Fig. 438. — Mouth-parts of larva of net-winged midge, Bibiocephala doanei. md., man- 
dible; mx., maxilla; l.ep., labrum-epipharynx; li., labium; hyp., hypopharynx. 
(Much enlarged.) 

theV occur in North America, South America, and Europe — entomologists 
believe that in past ages the family was much larger than it now is. 

The flies (Fig. 433) themselves can be distinguished when in hand by 
the curious secondary or pseudo net-veining of the wings. These faint cross 


The Two-winged Flies 

and diagonal veins are the marks of the creases made by the compact folding 
of the wings in the pupal shell. The females are provided with long saw- 
edged mandibles (Fig. 434), and are predatory in habit, catching smaller 
flying insects, especially Chironomid midges, lacerating their bodies with 
the mandibular saws and sucking the blood. The males have no mandibles, 
and probably take flower-nectar for food. Both males and females of several 
genera have the compound eyes divided 
into a large-facetted and a small-facetted 
part (Figs. 435 and 437). The egg-laying 
has not yet been observed, although the 
eggs must almost certainly be deposited 
on rocks in the stream or on its edge. 

With the mosquito wrigglers and the 
blood-worms (larvae of the Chironomidae) 
may perhaps be found a third kind of fly 
larva (Fig. 440), a slender, pale-colored, 
cylindrical Httle "worm," about one- 
third of an inch long, which can be 
distinguished from the other aquatic 
larvae by its two pairs of short leg-like 
processes borne on the under side of the 

Fig. 439. Fig. 440. 

Fig. 430. — Diagram of horizontal section through head of old larva of net-winged midge, 
Bibiocephala doanei, showing formation of adult head-parts inside,, larval 
mandible; /.wx., larval maxilla; /.c, larval cuticle; i.wrd., adult mandible;, 
adult maxillary palpus; id., hypoderm (cell-layer of adult skin of head); i.e., adult 
eye. (Much enlarged.) 

Fig. 440. — Larva of Dixa sp., with dorsal aspect of head in upper corner. (From life; 
much enlarged.) 

fourth and fifth body segments. It usually keeps the body bent almost double, 
and when feeding near the surface the head is twisted so that the under or 

The Two-winged Flies 


mouth side faces up although the rest of the body has its ventral aspect facing 
down. This larva belongs to one of the midge-Hke flies of the genus Dixa 
(Fig. 440), which is the only genus in the family Dixidae, represented by about 
a dozen North American species. The winged flies (Fig. 442) are found in 
moist places, densely grown over with bushes or rank herbage, in woods. 
Although resembling mosquitoes and 
Chironomid midges in general appear- 
ance, they can be readily distinguished 
from them by the arrangement of 
the wing-veins (Fig. 444). 

An interesting small group of 
readily recognizable flies is the 
family Psychodida% or "moth-fly" 
family. The vernacular name comes 
from the slight resemblance to minute 
moths shown by these flies because 
of the hairy broad wings, which are 
held over the back when the fly is at 
rest in the roof-like manner of the 
moths (Fig. 445). The largest of these Fig. 441. — ^Pupa of Dixa sp. 

flies are onlv about one-sixth of an -p^^^^f. '' n- 

riG. 442. — Dtxa sp. 

inch long, and are rarely distinguished 
except by careful observers. I have found them especially common in gar- 
dens near the seashore in CaUfornia, and also in the overhanging foliage 



(Much en-> 

(Much enlarged.) 

Fig. 443. — Mouth-parts of Dixa sp., female, l.ep., labrum-epipharynx; tnd., mandible; 
OTjf., maxilla; wx./., maxillary lobe; wjc./)., maxillary palpus; /i., labium; /»^., para- 
glossa; gl., glossa; hyp., hypopharynx. 

of trees and shrubs bordering the swift little mountain streams of the Coast 
Range. In one of these streams I was fortunate enough to find the 


The Two-winged Flies 

immature stages of one moth-fly species, Pericoma calijornica, which is, so 
far, the only North American member of this family whose life-history is 
known. The larvae (Fig. 446), which are Httle slug-hke creatures, one- 
tenth of an inch long, cling by a row of eight suckers on their ventral side 
to stones in or on the margin of the stream, where they are constantly 



Fig. 444. Fig. 445. 

444. — Diagram of wing of Dixa sp., showing venation. 

Fig. 445. — A moth-fly, Pericoma calijornica. (Much enlarged.) 

wetted by the dashing water. When ready to pupate the larvae crawl a little 
higher on the stones, where only the spray will reach them, and, fixing them- 
selves to the rock face by a gummy exudation, change to small flattish, 
turtle-backed pupae (Fig. 446), each with a pair of club- or trumpet-shaped 
respiratory horns on the back of the prothorax. They look indeed much 
like dwarf net-winged midge pupae. After ^ ^-i^i lmuPS^ 

about three weeks the adults issue and fly 



Fig. 446. Fig. 447. 

Fig. 446. — Larva, ventral surface (at left), and pupa, dorsal surface (at right), of the 
moth-fly, Pericoma calijornica; also enlarged prothoracic respiratory tube of pupa. 
(Much enlarged.) 

Fig. 447. — Mouth-parts of moth-fly, Psychoda sp. Ih., labrum; inx., maxilla; inx.p., 
maxillary palpus; mx.l., maxillary lobe; li., labium; pg., paraglossa; hyp., hypo- 

up into the overhanging foliage, where they spend most of their time 
resting on the under side of the leaves. 

The largest family of nematocerous flies in point of number of species, 

The Two-winged Flies 321 

and that one containing the largest flies in the whole order, is the family 
Tipulida;, whose long-legged, narrow-winged members are famiharly known as 
crane-flies, leather-jackets, and " granddaddy-long-legs. " The granddaddy- 
long-leg flies, which have wings, should not be confused with the often simi- 
larly named harvestmen, which are alhes of the spiders, have no wings, and 
have four instead of three pairs of legs. The Tipulid legs are ' extremely 
fragile, breaking off at a touch. Most slender-bodied, long- and thin-legged; 
two-winged insects of more than one-half-inch length of body are Tipulids. 
There are some smaller species, 
however, which might be mis- 
taken for midges or mos- 
quitoes, were it not that all 
Tipuhds bear a distinct V- 
shaped mark (suture) on the Fig. 448.— Diagram of wing of crane-fly, Sim- 

; . , , - ^ , plecta sp., showing venation, 

back of the thorax. More than 

three hundred species of this family are known in the United States, and they 
are common all over the country, in meadows, pastures, along roadsides, 
stream-banks, and in viroods. The flight is uneven, slow, and weak, and 
the ungainly fhes with their long middle and hind legs training out behind, 
and the front legs held angularly projecting in front, are unmistakable 
when seen in the air. 

The eggs are laid in the ground at the bases of grasses and pasture plants, 
or, by some species, in mud or slime. The footless, worm-like, dirty-white 
larvae feed on decaying vegetable matter, fungi, or on the roots or leaves of 
green plants. The root-feeders do some damage to meadows and pastures. 

The largest Tipulid, and the largest species in the whole order of flies, is 
the giant crane-fly, Holorusia ruhiginosa (Fig. 449), common in CaHfornia. 
Its body is nearly two inches long, and its legs are from two to two and one- 
half inches long, so that the spread of legs is four inches. The eggs are 
laid in the ooze of wet banks of little streams where faUen leaves are decay- 
ing and subdrainage water is always slowly trickling out from the soil. The 
larvae (Fig. 450) lie in this slimy bed, in crevices or on narrow ledges of rock, 
with the posterior tip of the body bearing the two breathing-openings (spi- 
racles) held at the surface. The soft ooze, composed of soil and slowly 
decomposing leaves, is swaUowed, and, as it passes through the ahmentary 
canal, the organic material digested out of it. The footless, worm-like 
larvae grow to be two and one-half inches long, but can contract to less than 
an inch. The duration of the larval life is not yet known, but it is at least 
several months. The pupae (Fig. 450), which are provided with a pair of 
long, slender respiratory horns on the prothorax, lie motionless in the slime 
for twelve days, when the great flies emerge and fly up into the foHage of 
the stream bank. 


The Two-winged Flies 

Next to the mosquitoes, the worst pests among the nematocerous flies are 
various species of the gall-midge family, Cecidomyidae, a family in which 
all the stages, larval, pupal, and adult, of all the species are terrestrial. The 

gall-midges are the frailest, 
smallest, and least conspicuous 
of all the flies, but their great 
numbers and vegetable feeding 

Fig. 449. Fig. 450, 

Fig. 449. — The giant crane-fly, Holorusia rubiginosa, male. (Three-fourths natural 

Fig. 450. — Larva (at left) and pupa (at right) of giant crane-fly, Holorusia rubiginosa; 

in middle of figure enlarged posterior aspect of larval body, showing spiracles. 

(Larva and pupa three-fourths natural size.) 

and gall-making habits make them formidable enemies of many of our 
cultivated plants. The tremendous aggregate losses suffered by the wheat- 
growers of this country from the ravages of the Hessian fly, the damage 
to clover-fields by the clover-leaf and clover-seed midges, and the injuring 
, or killing of thousands of pine-trees from the attacks of the minute 
pine Diplosids, are evidences of the great economic importance of the 
delicate little gall-gnats. About one hundred species are known in this 
country, and of these most are more or less destructive to cultivated herbs, 
shrubs, or trees. 

The tiny bodies of the flies are usually covered with fine hair, easily 
rubbed off, and the antennae bear whorls of larger hairs, which, with some 
species, are attached by both ends, thus making little hair loops. The 
minute eggs, reddish or white, are usually deposited in or on growing plant- 
tissue, and the little footless, headless, maggot-like larvae probably derive 
most of their food by imbibing it through the skin. Lying with the body 

The Two-winged Flies 323 

practically immersed in plant-sap, the thin body-wall acts as an osmotic 
membrane through which an interchange of fluids takes place automati- 
cally. The Cecid larva has to eat whether it will or not, and has to eat 
practically all of the time! These larvai may be distinguished by their 
possession of a strange little cliitin plate on the under side of the front part 
of the body, called the breast-bone. What the exact use of this little sclerite 
is has not yet been determined. Perhaps it helps in locomotion, perhaps 
in rasping or lacerating the soft plant-tissue to increase the flow of sap. The 
larvae pupate where they lie, sometimes spinning a thin silken cocoon, some- 
times transforming within the hardened last larval moult, sometimes with 
no special protecting covering at all. 

The most notorious gall-gnat is the wheat-pest, known as the Hessian 
fly, Cecidomyia destructor, and distributed over all the United States east of 
meridian 100°, as well as in California. By the ravages of its larvae, feeding 
as they do on the sap of growing wheat, this minute fly causes an annual loss 
in this country of approximately ten million dollars. This enormous direct 
tax is paid by those farmers who prefer to farm in the good old way, with a 
strong belief in the dispensations of an erratic Providence, rather than to 
do their farming as modified by modern knowledge and practice. The 
tax-collecting insect, which is a tiny delicate blackish midge about one- 
tenth of an inch long, lays its eggs in the creases or furrows of the upper 
surface of the leaves of young wheat, and the hatching larva; wriggle down 
to the sheathing bases of the leaves, where they lie and drain away the sap 
of the growing plant. When full-grown they pupate within the outer hardened 
brown last larval cuticle, and resemble very much a, small spindle-shaped 
seed. This is called commonly the "flaxseed" stage. The adult soon 
issues and after a few days of flight and egg-laying dies. There may be as 
many as four or five generations in a year, both spring and winter wheat 
being attacked. The remedies are the late planting of winter wheat, the 
burning or plowing in of the stubble after harvesting, and the early planting 
of strips of decoy wheat about the field, which shall attract the egg-laying 
females and may be afterwards plowed under with the myriad eggs it contains. 
The Hessian fly is a European insect brought unintentionally to this country 
about 1778, but probably not, as often said, with the straw brought by the 
Hessian troopers of the Revolutionary War. It attacks rye and barley as 
well as wheat, and has, in turn, to withstand the combined attacks of half 
a dozen hymenopterous parasites, which are said to destroy nine-tenths 
of all the Hessian-fly larvae. Without these natural checks to its increase 
this pest would destroy every wheat-field in this country in a very few 

In 1896 the Monterey pines, Pinus radiata, much grown, together 
with the famous Monterey cypresses, as ornamental trees on the San Fran- 


The Two-winged Flies 

Cisco peninsula, showed a peculiar stunting and gall-like swelling of the 
leaves. Since then this deformation has appeared so abundantly and widely 
within the range of this tree that the species is actually threatened with 
extinction, the shortened, swollen needles not being able to perform the 
essential food-assimilating functions of green leaves. This injury is due 
to a single species of Cecid fly known as Diplosis pini-radiata (Fig. 451), 

Fig. 451. — The Monterey-pine midge, Diplosis pini-radiata; eggs in upper left-hand 
corner; pupa, larva, breast-bone of larva, and adult female. (Much enlarged.) 

which lays its eggs at the base of the growing new needles and whose larvae 
hatching and lying here use up the sap necessary for the development of 
the needles. Hundreds of Monterey pines have been cut down, and unless 
the natural enemies of this little fly, of which two or three have been dis- 
covered, get the upper hand of the pest, this splendid species of pine may 
be wholly destroyed. A half-dozen other species of Diplosis are known 
in this country and Europe as pests of conifers, but no other pine species 
seems to have suffered quite so severely as this interesting Cahfornian one, 
whose whole geographical range extends over but a thousand square miles, 
and which is thus specially liable to destruction by concentrated insect 

If the collector will break up and examine carefully almost any old or 
partially decaying toadstools or shelf fungi from trees, he will find in the 
soft fungous body numerous small translucent white maggot-like larvae, the 
larvae of fungus gnats or members of the family Mycetophilidae. The gnats 
themselves are slender delicate flies, mostly with clear wings, though some 
common species have dark wings, with the basal segment (coxa) of the legs 
unusually long and the antennae in most cases free from the whorls of long 
hairs so characteristic of the Chironomidae, Culicidae, and other families of 
flies otherwise much resembling the fungus-gnats. The flies are to be looked 
for on decaying vegetable matter, especially fungi, and in damp places. 

The eggs are laid variously: on fungi, in decaying wood, among decom- 
posing leaves, in animal excrement, and under the bark of trees. The larvae 

The Two-winged Flies 


feed on the decomposing substance in which the eggs are ^aid, sometimes 
spinning silken webs for protection. They pupate in the food-substance or 
crawl away to some more sheltered spot, often forming a thick cocoon in 

Fig. 452. — A fungus-gnat of the family Mycetophilida; larva, pupa, and adult. 

(Much enlarged.) 

which to transform. Perhaps the most singular habits noted in the family 
are those connected with the strong gregarious instinct which leads the 
larv£e of many species to hve closely together. Some of the species of Sciara, 
known as "army-worms," have "the singular propensity of sticking to- 
gether in dense patches, and will 
form processions sometimes twelve 
or fourteen feet in length and two 
or three inches broad. This phe- 
nomenon has been observed fre- 
quently both in Europe and Amer- 
ica, but the reason therefor is not 
yet well understood, though the 
object of the migration seems to be 
the search for better feeding-grounds." Various species of this genus live 
in potatoes and other vegetables, while the serious injury to potatoes called 
"scab" is caused by a fungus-gnat known as Epidapus scabies. 

With larger and more robust bodies and relatively shorter and thicker an- 
tenncT, the March-flies, Bibionida?, serve as a sort of transition family between 
the long-legged, slender -bodied midge type of fly with its thread-like hairy 
antennae, and the compact, heavy-bodied, short-legged type of fly with short 
and club-Hke three-segmented antennae, characteristic of the many families 
grouped in the section Brachycera. The March-flies (Fig. 454) are from 
one-eighth to one-half inch long, with fairly robust, often hairy, body, black- 

FiG. 453. — Diagram of wing of fungus-gnat, 
Mycetophila sp., showing venation. 


The Two-winged Flies 

ish or blacK and red, strong legs, large clear or smoky wings, and stout an- 
tennae about as long as head and thorax together and composed of nine to 
twelve segments. They may be seen often in large numbers flying heavily 
over gardens and fields or in woods, early in the spring. The eggs are laid 
in the soil or in decaying vegetation or in sewers and excrement, the larvae 
feeding usually on decom.posing substances. With some species, however, 
the larvas feed on the roots of grains or grasses and in this way may do serious 
damage. Bibio tristis, discovered in Kansas in 1891, appeared in great 
numbers in wheat-fields and frightened many wheat-growers. As a matter 
of fact, Httle injury seemed to be done. B. jemorata, a common species, 
is deep red with black wings; B. albipennis, another abundant and wide- 
spread one, is black-bodied with white wings. A common CaHfornian species 
appears from the ground in damp woods in great numbers in March. I 
have watched these flies issuing in countless numbers 
from the soft rich forest floor in the extensive 
Monterey pine woods near the Bay of Monterey 

Fig. 454. Fig. 455. 

Fig. 454. — March-fly, Bibio albipennis. (Three times natural size.) 
Fig. 455. — Diagram of wing of Bibio albipennis, showing venation. 

The air danced with them, and the pine-trees and shrubs bore countless 
myriads on their branches. Professor Needham records a similar sight 
in which individuals of B. jraternus formed the hosts, and a woodland pasture 
near Lake Michigan was the scene of their appearance. "I have rarely 
come upon a scene of greater animation than a sheltered hollow in this wood 
presented," writes Professor Needham. "There was the undulating field 
clad in waving grass and set about with the pale-hued foliage of the white 
oaks; there were the flowering hawthorns; and there were the myriads 
of Bibios floating in the sunshine, streaming here and there like chaff before 
sudden gusts and swirls of air. All the spiders' webs in the bushes were 
filled with captives; Httle groups of ants were dragging single flies away to 
their nests, and once I saw overhead a chestnut-sided warbler, perched on 
a bare bough directly in a stream of passing flies, rapidly pecking to right 
and to left, persistently stuffing his already rotund maw. I counted a number 
of flies I could see resting on the grass in several small areas wide apart, and 

The Two-winged Flies .327 

found the counts averaged fifteen Bibios per square foot; and there were 

here in one place forty acres of such Bibio territory." 

Two famihes of nematocerous flies are not included in the key, and have 

not heretofore been referred to. They are the Orphnephihdae, of which but 

a single species is known in this country, viz., Orphnephila testacea, a small 

reddish-yellow fly without hairs or bristles on its body, and with short antennae 

apparently composed of two segments, but really of ten, the apparent first 

segment being made up of three closely 

opposed segments, and the second of seven. 

The fly itself is found along stream banks, 

but nothing is known of its immature stages. 

The other family, Rhvphidae, or false crane- ^ 

.... . 1 ^ 1*10. 456. — Diagram of wing 

flies, is represented in this country by two Rhyphus sp. 

genera containing several species. The flies 

are small and slender, with broad spotted wings veined in a character- 
istic way (Fig. 456). The larvas of Rhyphus are worm -like, legless, naked, 
more or less transparent, with snake-like movements. They live in water, 
brooks, pools, or puddles, or in rotting wood, hollow trees, or manure. 


The Brachycera, or flies with "short horns," i.e., short thick antennae 
composed of few segments, in contrast with the many-segmented antennae, 
usually slender and long, of the Nematocera, are separable into three groups 
of famihes, as indicated in the key on page 303, based on a further analysis 
of the structural character of the antennae. These groups are, first, one includ- 
ing flies in which the antennae are composed of more than five segments but 
with all those beyond the second coalesced to form a single compound 
segment, bearing more or less distinct annulations indicating the component 
subsegments; second, one including flies having antennae made of four or 
five distinct segments; and third, and by far the largest, one including flies 
with but three segments in the antennae. 

In the first group are two families and part of a third; this division of a 
family indicating plainly the artificial character of the subdivision into 
groups, the subdivision being merely convenient. The three famihes may be 
distinguished as follows: 

The branches of the radial vein (see Fig. 460) crowded together near the costal (front) 

margin of the wing (Soldier-flies.) Stratiomyid.*;. 

Venation normal. 

Alulets, i.e., little whitish wing-like membranous flaps at the base of the true wings, 

large (Horse-flies.) Tabanid^. 

Alulets small (Snipe-flies.) Leptid^ (in part). 


The Two-winged Flies 

The most familiar and interesting flies in this group are the well-known 
horse-flies, gad-flies, or deer-flies, Tabanidas. They are all fairly large, 
some indeed being among the largest of our flies. 

The great, black, swift horse-flies that in summer dart suddenly at our 
carriage-horses and with quick shifting flight seem to be fairly carried 
along in the air close to the horses, are the most familiar representatives of 

Fig. 457. — Greenhead, or horse-fly, Tabanus lineola. 

indicated by line.) 

(After Lugger; natural size 

the order. Many of the smaller horse-flies show gleaming metalHc colors, 
especially about the head. Much of this color is in the large compound 
eyes, and almost any horse-fly caught alive or just killed will astonish the 
collector by the brilliant bands and flecks of iridescent green, violet, purple. 

Fig. 458. — Diagram of wing ot Chrysops sp., a horse-fly, showing venation. 

and copper on the eyes. The biting and blood-sucking are done by the 
females alone, the males lacking the sharp dagger-like piercing mandibles 
and contenting themselves with lapping up flower-nectar. 

The brown elongate eggs of horse-flies are laid either on stems or leaves 
of terrestrial plants, or on aquatic plants or submerged stones. The larvae, 
whitish, cylindrical, tapering at both ends, and with a series of slightly raised 
roughened ridges running around the body, either live in water, in slimy 
places along pond and brook shores, or in soft rich soil, and are predaceous, 

The Two-winged Flies 


feeding on small aquatic or underground creatures, especially insect larva 
and snails or slugs. 

Nearly 200 species of horse-flies are known in North America. The 
large bluish-black and brownish-black ones, an inch long and with dusty 
wings expanding for two inches or more, belong to the genera Tabanus and 
Therioplectes; the smaller "greenheads" with banded wings and briUiantly 

^'''■«J^/~^°'^n^"P''ri°^ ^ horse-fly, Therioplectes sp. md., mandible; mx., maxilla; 
mx.l., maxillary \ohy mx.p., maxillary palpus; hyp., hypopharynx; lb., labrum 
ep., epipharynx; h., labium; la., labellum. ^ -'i' 1 ^ > . 

colored eyes and black or brown and yellow bodies mostly belong to the 
genus Chrysops. Silvius pollinosiis is a beautiful small species with a milk- 
white bloom over its body, and with clear whitish wings with a few small 
brown spots. 

The soldier-flies, Stratiomyidae, are unfamiliar insects, although as many 
species of them as of horse-flies occur in this country. Many of the species 
have bright yellow or green markings, and most of them have the abdomen 
curiously broad and flattened. 
They are found about flowers, 
and can readily be classified, 
after capture, by the unusual 
character of the venation (see 
Fig. 460). The eggs are laid 
on the ground or on ^leaves in or 
near water, some of the larvc-e 
being terrestrial, while others are 
aquatic. The food seems to be mostly vegetable, although the larva of some 
species are believed to be carnivorous. One or two species live in salt or 
brackish water, and Sharp records that some Stratiomyid larva were found 
in a hot spring in Wyoming with the water temperature only 20° to 30° F. 
below boihng. They pupate within the last larval skin, which is long and 

Fig. 460. — Diagram of wing of Odontomyia 
sp., showing venation. 

33° The Two-winged Flies 

tapering at one end. Some species inhabit ants' nests, and one is suspected 
of living parasitically in bee-hives. 

Stratiomyia is a genus containing rather large conspicuous yellow-banded 
flies with broad flattened abdomen, while Sargus, a genus whose species 
are common, has a subcyhndrical abdomen with the whole body metallic 

The snipe-flies, Leptidae, are a small family represented by about fifty 
North American species, including flies having no habits or structural pecu- 
liarities appealing specially to popular interest. They are rather slender 
and plainly colored, and rather heavy and slow in movement. They are 

apparently all predatory in both larval 

and adult stages. The adults may be 

best found, according to Comstock, in 

low bushes and grass. The larvae Hve 

in the ground, in moss, or in decaying 

wood, sometimes penetrating to the 

Fig. 461.— Diagram of wing oi Chryso- burrows of wood-boring insects. The 

phila thoracica (LeptiQEe), showing . k ^ • ^ 

venation. species of the genus Atherix deposit 

their eggs "in dense masses attached 
to dry branches overhanging water. Not only do numerous females con- 
tribute to the formation of these masses, but they lemain there themselves 
and die. The larvae on hatching escape into the water." 

In the second group of Brachycera, including flies which have their anten- 
nae composed of four or five distinct segments, there are two famihes, the 
Asilidae, or robber-flies, and the Midaidae, or Midas-flies. These latter resemble 
the robber-flies in size and general appearance, but differ from them by having 
the antennae rather long and clubbed at the tip. They are predaceous, 
catching and devouring other flying insects, and the larvae of the few species 
whose life-history is known are also carnivorous, and seem to have a special 
fancy for the larvae of the great wood-boring grubs of the giant Prionus 
beetles. Howard believes that the large species, Mydas luleipennis, found 
in the Southwest, mimics in coloration and general appearance for protection 
or aggression the tarantula-killer wasp found commonly in this country. 

The Asilidae, or robber-flies, compose a considerable family — nearly 1000 
species occur in this country — of large, swift, hairy, ferocious-looking flies 
which hve wholly by predatory attacks on other insects. The body is usually 
long and slender, tapering behind (Fig. 462), although in a few genera the 
abdomen is flattened and not unusually elongate. The proboscis is strong 
and sharp, the eyes large and keen, and the wings long and narrow and 
capable of carrying this insect hawk swiftly and strongly in pursuit of its 
prey. Some of the robber-flies are very large, an inch and a half or even 
two inches long, and they do not hesitate to attack other large and strong and 

The Two-winged Flies 


well-defended insects, as bumble-bees, dragon-flies, and the fierce and 
active tiger-beetles. The robber-flies usually rest on the ground or on low 

Fig. 462. Fig. 463. 

Fig. 462. — A robber-fly, Stenopogon inquinatus. (Natural size.) 
Fig. 463. — A bumble-bee-like robber-fly, Dasyliis soceata. (Natural size.) 

foliage, and fly quickly up with a buzzing sound when disturbed or attracted 
by prey. All the prey is caught on the wing, held in the long spiny feet of 
the robber-fly, and torn and sucked dry by the sharp piercing-beak. 

Fig. 464. — ^Diagram of wing of robber-fly, Erax cinerascens, showing venation. 

The larvae live chiefly in decaying wood or in soil containing decom- 
posing vegetable matter, and are also predatory, feeding on grubs and other 

Fig. 465. — Mouth-parts of robber-fly, Erax cinerascens. li., labium; hyp., hypopharynx; 
lb., labrum; mx., maxilla; mxL, maxillary lobe; mxp., maxillary palpus. 

underground or wood-boring insects. The pupje are curiously spiny, the 
spines being used as a sort of pushing or pulling organ when they get ready 
to come to the surface of the ground or dead tree to change into imagines 

Some of the species of the genera Laphria and Dasyliis (Fig. 463) look 
astonishingly like bumble-bees and wasps, probably a case of protective 

332 The Two-winged Flies 

mimicry (see Chap. XVII). Erax is a gemis with many common gray and 
black species about an inch long, with sharp-pointed tip of the abdomen. 

The third section or group of Brachycerous famihes includes many 
families, in all of which the antennae have the first two segments small and 
the third curiously large and club-like, and usually bearing a single con- 
spicuous bristle-like hair. The families of this group can be distinguished 
by the following table: 

A. Antennae composed of three segments, the third usually large and either with or 
without a bristle or style. 
B. Empodium pulvilliform, i.e., feet with three little pads instead of two. 

(Snipe-flies.) LEPXiDiE (in part). 
BB. Empodium not pulvilliform, i.e., feet with two little pads and a median bristle 
or nothing. 
C. Radial vein four-branched. 

D. Second branch of cubital vein extending free to the margin of the 
wing or coalesced with the first anal vein for a short distance 

(see Fig. 466) (Bee-flies.) Bombyliid^. 

DD. Second branch of cubital vein joining first anal far from the 
margin of the wing (see lig. 471). 

(Dance-flies.) Empidid^ (in part). 
CC. Radial vein with not more than three branches. 

D. Head with a curving suture immediately above the antennas. 

(House-flies and allies.) MusciD^. 
DD. Head without such suture. 

E. Radial vein with a knot-shaped swelling at the point where 
it forks, with a small cross-vein running back iust at or near 
this sweUing (Fig. 474). .(Long-legged flies.) Dolichopodid^. 
EE. Wings without such characteristics. 

F. Second branch of cubital vem appearing as a cross- 
vein or curved back towards the base of the wings 
(Fig. 479). 

G. Proboscis rudimentary; mouth-opening small; palpi 
wanting; antennae with dorsal arista. 

(Bot-flies.) CEsTRiD^. 
GG. Proboscis not rudimentary; palpi present; antennae 
with terminal style or arista or dorsal arista. 

Empidid^ (in part). 
FF. Second branch of cubital vein not appearing like a 
G. Front with grooves or a depression beneath the 

antennae (Wasp-flies.) CoNOPiDiE. 

GG. Front convex beneath the antennae; a spurious 
vein usually present between radius and media 
(Fig. 479) (Flower-flies.) SyRPHiD.E. 

The famihes of flies named in the above key contain many hundreds of 
species but few of which are at all popularly known. The bot-flies (CEstridae), 
house-flies, flesh-flies, bluebottles and stable-flies (Muscidas calyptrata), and 

The Two-winged Flies 333 

the cheese-skippers and pomace-flies (Muscidae acalyptratae) are about the 
only names in the Hst of these hundreds which seem at all familiar. The 
flower-flies (Syrphida?) and bee-flies (BombyHidaj) are numerous, often 
seen, and, what is more, often definitely noted and admired, but "beautiful 
flies" is about as specific a name as they ever get. The bristly parasitic 
Tachinid flies are noticed now and then by the nature student, and the 
dancing Empidids interest, in a decided but irritating way, drivers and 
bicyclers in the dance-fly mating-time. But even entomologists, professional 
as well as amateur, unless they are special collectors and students of 
Diptera, recognize but few of the hosts of small flies that fill the air during 
the long summer days. 

In the above key only the larger and more commonly represented famihes 
are included, so that it will be possible for a collector using this book to 
find himself possessed of a fly which will prove intractable when an attempt 
is made to classify it into its proper family. But such unfortunate happen- 
ings will be very infrequent, as only small families of obscure or rare species 
are thus omitted. 

Poised almost motionless in the air a few inches above a sunny path or 

roadway, or darting away, when disturbed, with lightning swiftness and 

having all the seeming of bees, hairy, plump-bodied, and amber-colored, certain 

bee-flies (Bombyhidas) are rather familiar acquaintances of the summer field 

student. Other bee-flies, as swift 

and as beautiful, are less bee-like 

because of the striking "pictures" 

in the wings, blackish or brown 

blotches conspicuous in the thin, 

otherwise clear wing-membrane. 

Some of these bee-flies have ' an 

1, , , J , . Fig. 466. — Diagram of wing of Anthrax ful- 

unusually long slender proboscis ^/a,°, showing venation. 

held straight out in front of the 

head like a spear at rest (Fig. 467). But this beak has no bloodthirstiness; it 
is used to suck up sweet nectar from flower-cups. The larvae of the bee-flies, 
however, are carnivorous, Hving parasitically in the egg-cases of grasshoppers 
or on the bodies of wild bees and various caterpillars. One of these bee- 
fly larvas burrowing into a grasshopper's egg-pod can do awful harm to the 
embryo grasshoppers, but at the same time much good to us, by the satisfac- 
tion of its egg-eating propensities. Beautiful, velvet-clothed, swift-winged, 
and nectar-feeding as a fly, maggot-Hke and parasitic as larva, the bee-fly 
is a good example of the great differences in structure and habit which are 
possible between young and old of the speciaHzed insects. 

Bombylius (Fig. 467) is a genus in which the proboscis is very long and 
slender, the body short and plump and covered with a thick soft coat of longisb 


The Two-winged Plies 

hair usually light brown or whitish in color. The wings are blotched with 
brown or blackish. Anthrax contains numerous species with short proboscis, 
and broad flattened body covered with short hair. The wings are either 
clear or partly colored with brown or black. In the species of the genus 
Exoprosopa (Fig. 468) the hair of the body is very short and often in silvery 

bands across the abdomen, the pro- 
boscis is short, and the wings usually 
beautifully "pictured" with brown and 

Fig. 467. 
Fig. 467. — A bee-fly, Bombylius major 
Fig. 468. — A bee-fly, Exoprosopa sp 

Fig. 468. 
(Twice natural size.) 
(One and one-half times natural size.) 

In California the roads and paths, especially along streams and through 
woods and parks, are made almost intolerable in part of the spring for driving 
or bicychng because of hosts of small slender blackish flies 
in swiftly dancing swarms. These are dance - flies, 
Empididas, and their aerial dance is their mating flight. 
I do not know that such hordes of dance-flies occur in 
the East, but some species of 
the family have the same danc- 
ing habit there, and can be dis- 
tinguished by it and by the 
structural characters given in 
the key. The midges, Chirono- 
midae, also dance in swarms in 
the air, but are readily dis- 
tinguished from the Empidids 
by their small fragile body, 
and long many-segmented hairy 
antennae. All the dance -flies 
are predaceous, sometimes 
catching their prey in the air, 
sometimes chasing it on the 
ground. The larvae, slender cylindrical grubs living in the soil or under leaves 

Fig. 469. Fig. 470. 

Fig. 469. — Mouth-parts of a bee-flv Bombylius sp 

(Much enlarged.) 
Fig. 470. — A dance-fly, Rhamphomyia longicauda 

(Three times natural size.) 

The Two-winged Flies nor 

or other vegetable matter, are also probably predaceous, feeding on smaller 
insects living in the same places. 

The commoner species that dance in large swarms belong to the genera 
Empis and Rhamphomyia (Fig. 470). The males of certain species of Empis 
and Hilara have the odd habit of blowing out bubbles of a whitish viscid sub- 
stance which they carry about with them in the air. It is believed that these 
toy balloons are attractive to the females. At least, Professor Aldrich, a 
well-known student of flies, has seen a female choose that male among several 
which was carrying the largest balloon! 

An attractive lot of small slender flies, usually of iridescent green or 
greenish-black or blue color, with 
unusually long slender legs, are 
the Dohchopodidas, or long-legged 
flies. They are found especially 
in marshy or low places where 
vegetation grows lush and rank. 
They flit about searching for 
lesser insects, which they catch 
and devour. They often get their 

• 471- — Diagram of wing of dance-fly, 
Rhamphomyia sp., showing venation. 

prey by swift chasing over leaves or ground or even on the surface of water. 
Like the Empidids the larvas are also predaceous, living underground or in 
decaying vegetable matter. Some have been found in the exuding sap of 




Fig. 472. Fij, ^^^ 

472.— Mouth-parts of dance-fly, Rhamphomyia sp. lb., labrum; \,!x., maxilla; 
Ftp "!^; ' "^^ r ^"I ""Yi f ^•/'•' "^^^^iHary palpus; li., labium; hyp., hypopharynx. 
I'lG- 473-— Dohchopus lobatus. (Three times natural size.) 

trees and elsewhere on or under bark. The larvae of certain species spin 
little thin cocoons when ready to pupate, but with most the pupa is 


The Two-winged Flies 

Dolichopus (Fig. 473) is the largest genus of the family, nearly 100 species 
occurring in this country. The males are curiously ornamented by special 
outgrowths or expansions on the feet. These make the feet at the end of 
the long legs very conspicuous and are believed to serve the male to help 
attract the female in his courtship of her. These ornaments are not con- 
fined to the males of this genus, other genera of the family showing similar 

Fig. 474. — Diagram of wing of a Dolichopodid, Psilopics ciliatus, showing venation. 

characters. Other ornaments, too, are found in various species, some occur- 
ring on the face, others on the antennae and elsewhere. Aldrich says that 
the males of the flies of this family show more pronounced and various special 
ornamentation than the males of any other single family of animals. He 
has seen the males dangle their tufted feet in the faces of the females during 

Occasionally the general collector or nature observer will find an insect 
that he has taken at first glance for a wasp, but which on examination, after 
capture, is found to have but a single pair of wings, and short, clubbed anten- 
na; like a fly. The puzzle is readily solved with 
these clues: the insect is a fly, not a wasp; it simply 
looks so much hke a wasp that it undoubtedly is 
frequently mistaken for a wasp by certain enemies 
which are afraid to attack the well-defended hornet, 
but would make short work of a defenceless fly. 
The wasp-flies, Conopida;, thus save their lives by 
an innocent deception; they are protected by their 
curiously close mimicry of wasps. All of them are 
narrow-waisted, and most have the abdomen spindle- 
shaped and tapering like a wasp's, and often banded 
and colored so as to increase the similitude. All 
of them, too, have robust heads and have been sometimes called "thick- 
head-flies." They are all flower-flies, feeding on nectar and pollen, and 
hovering on heavy wing about blossoming shrubs. The oval or pear-shaped 
larvje are parasitic, living in the bodies of other insects, especially wasps, 

Fig. 475. — A wasp-like 
Qy,PhysocephaIa affinis. 
(One and one-half times 
natural size.) 

The-Two-winged Flies 007 

bumble-bees, and locusts. "The eggs," according to Williston, "are laid 
directly upon the bodies of the bees or wasps during flight. The young 
larvae burrow within the abdominal cavity of their host and there remain, 
the posterior end directed toward the base of the abdomen, feeding upon 
the non-vital portions, until ready to transform into the mature fly, when they 
escape from between the abdominal wings of the insect.'' The quiescent 
pupal stage is then passed within the body of the host, a rather unusual 
phenomenon in insect Hfe. 

In the genera Conops and Physocephala (Fig. 475) the abdomen is distinctly 
peduncled as in the thread-waisted wasps, while in Myopa, Zodion, Oncomyia, 
and others the abdomen is sessile or constricted only at the very base. 

Under the name bot-flies (CEstridae) some of the most interesting members 
of the order Diptera are widely, but superficially, known. The flies themselves 
are much less familiar than their eggs and larvae, the glistening white eggs 
of some species being often seen attached to the flanks, legs, 
or feet of a horse or cow, and the stomach-inhabiting larvoe 
being well known to stockmen as the cause of much suffer- 
ing and injury to their animals. In addition to the "bots" 
which live in the stomach and intestines of horses and 
cattle, several other species live under the skin of the same 
animals, as well as of goats, sheep, antelope, rabbits, rats. Fig. 476. — Larva 

dogs, cats, and even man. The larvae of still other species ? "°^""7' yj''^''^- 
° ' ' ^ bra cimtculi, from 

burrow in the nasal passages of the sheep, the antelope, wood-rat, Neoto- 
the horse, the camel, the buffalo, and various deer species, "f*^ ^P- (Natural 
The flies are heavy-bodied, often densely hairy, banded in- 
sects, looking rather like small bumble-bees whose mouth-parts are so atrophied 
that they can probably take no food at all. They lay their eggs on the hairs 
or skin of their special host animal, and the larvae on hatching bore directly 
through the skin and into the tissues of the host, or, as in the case of the 
familiar bot-fly of the horse and the heel-fly or warble of cattle, the eggs are 
taken into the mouth of the host by hcking, swallowed, and thus introduced 
directly into the stomach, to whose walls the larvae either attach themselves or 
through which they burrow into the true body-cavity of the host. 

Less than 100 species of bot-flies are known in the whole world, 
but the parasitic habits and resulting economic importance of these flies 
have resulted in making the family well known. The most widely dis- 
tributed and best known species is probably the horse bot-fly, Gastrophiliis 
equi (Fig. 477). This fly, which may be seen in open sunny places along 
the roadways, is about ^ inch long, brownish yellow, with some darker 
markings, but much resembling a honey-bee in appearance. The female 
has the abdomen elongate and bent forward underneath the body. The 
light-yellow eggs are attached by a sticky fluid to the hair of the horse 


The Two-winged Flies 

on the shoulders or legs or belly. They are licked off by the horse and 
swallowed, and the larvae hatch in the mouth or stomach and attach themselves 
to the stomach lining, living at the expense of the host. When many larvae 
thus live in the stomach (and as many as several hundred have been found 
in one animal) the horse suffers serious injury. The larvae live in the stomach 

Fig. 477.- 

-Bot-fly of horse, male, Gastrophilus eqiii, abdomen of female and egg. 
Lugger; natural size of fly indicated by line.) 


and intestines through fall and winter, and late in the spring release their 
hold, pass through the intestine with the excretions, and burrow into the 
ground to pupate. The pupal stage lasts about a month, when the flies 
issue and the life-cycle begins again. A smaller species of bot-fly, Gastro- 
philus nasahs, with bright-yellow band across the abdomen, lays its eggs 
in the lips and nostrils of horses. For the rest its life-history is about like 
that of G. equi. 

The bot-flies, warble-flies, or heel-flies of cattle, whose larvae are found in 
small tumors under the skin, also have their eggs swallowed, and the young 
larvae may be found in the mouth and oesophagus. But from here they burrow 
out into the body-tissues of the host, finally coming to rest underneath the 
skin along the back. When the larva or grub is full-grown it gnaws through 
the skin, drops to the ground, pupates, and in from three to six weeks changes 
to the adult fly. The hides of cattle attacked by these flies are rendered 
nearly valueless by the holes, and are known as "grubby" hides. Osborn 
estimates that these warble-flies, of which we have two species, Hypoderma 
bovis and H. lineata, cause a loss of $50,000,000 annually in this country. 

The genus Cuterebra includes a number of species of which the rabbit 
bot-fly, C. cuniculi, is most familiar. The larvae lie in large warbles or tumors 
under the skin of the infested rabbit, and late in the summer the jack-rabbits 
and cottontails are so badly infested in some localities that hardly one can 
be found free from the pest. The adult is a large fly resembling a bumble- 

The Two-winged Flies 339 

bee, with black head, yellow-brown thorax, and the abdomen blue-black 
with yellow base. The full-grown larva is a large black spiny grub. 

One or two species of bot-flies infest man, and also (probably the same 
species) monkeys and dogs and perhaps other animals. Numerous instances 
are recorded in which the larvae of Dermatobia noxialis and D. cyaniventris 
have been found under the skin of persons in tropical America, and a few 
instances of such cases in the United States. The larvae are thick and broad 
at one extremity and elongate and tapering at the other. 

The family SyrphidiE, Syrphus-flies, flower-flies, or hover-flies, as the 
Enghsh call them, is one of the largest in the order; including fully 2500 
species in the whole world, of which over 300 are found in this country. 
For so large a family few generalizations regarding the appearance or 
habits of the flies can be made. Many of the Syrphus-flies resemble bees 
and wasps in appearance, and almost all are rather bright and handsome 
insects. They feed on nectar and pollen, and hence are to be found in sun- 
shiny hours at flowers, hovering like tiny humming-birds in front of open 

Fig. 478. Fig. 479. 

Fig. 478. — A flower-fly, Eristalis tenax. (One and one-half times natural size.) 
Fig. 479. — Diagram of wing of Syrphus continuax, showing venation. 

blossoms, or crawling bee-like in and out of deep flower-cups. Some make 
a distinct humming or buzzing as they fly about and thus heighten their 
suggestion of bees. All can be distinguished, after capture, by the so-called 
false vein of the wings (see Fig. 479). The larva? live variously in decaying 
wood or other vegetation, or decomposing flesh, or in the stems of green 
plants, or in toadstools, or in water. Some crawl about, slug-like in manner, 
over leaves, preying on aphids and scale-insects. Some live as guests in ants' 
nests, and others in the underground nests of bumble-bees. 

Those Syrphid larvae most often written about are the curious "rat-tailed 
maggots" (Fig. 480), larvae which live in stagnant water or slime and have 
the posterior extremity of the body greatly elongate and projecting to serve 
as a breathing-tube. There is a spiracle (breathing-pore) at the tip of this 
"tail," and the tail projects upward so that its tip reaches the air, while the 
rest of the larva's body remains underneath the water. The larvae of Micro- 


The Two-winged Flies 

don, which live in ants' nests, look like little mollu^s, and when first found 
were actually described as new molluscous genera. Their body is fiat. 

Fig. 480. Fig. 481. 

Fig. 480. — Rat-tailed larva of a Syrphid. (Twice natural size.) 
Fig. 481. — Larva of Microdon miUabilis, dorsal view. (Four times natural size.) 

broad, unsegmented, and looks Hke a fiat broadly elliptical little shell or 
plant-seed (Fig. 481). 

Among the more common flies of this family which may be taken by the 
collector are various species of Eristalis, with black, yellow, and amber colors, 
heavy-bodied, bee-like forms, and especially E. tenax, the drone-fly, which 
resembles very much a honey-bee drone. Its larva is a rat-tailed maggot. 
The species of Syrphus are black with yellow bands, with the abdomen 
not so heavy as in Eristalis. The larvae are predatory, doing great havoc 
in aphid colonies, but being thus of great benefit to florists and gardeners. 

Fig. 482. 

-Mouth-parts of Eristalis sp. li., labium; hyp., hypopharynx; Ih., labrum; 
mx., maxilla; mx.l., maxillary lobe; mx.p., maxillary palpus. 

The species of Volucella are bee-like in appearance and their larvae live in 
the nests of bees, but whether as parasites or tolerated guests seems not 
to be yet known. Sharp thinks that they act as scavengers in the nests, 
and thus are helpful rather than harmful to their hosts. Syritta pipiens is 
a common Syrphid fly, with slender, elongate, subcyUndrical body, blackish 
with reddish-yellow markings. 

The Two-winged Flies 


The abundant house-flies are the most famihar representatives of the 
largest of all the Dipterous families: largest if the great heterogeneous 
group of flies called Muscidae is to be looked on as a single family, a point of 
view taken by some entomologists, but not so if this group is called a 
superfamily^ composed of a large number, about twenty in all, of distinct 
small families. The group includes, besides the house-flies, the buzzing 
bluebottles, the disgusting flesh-flies and stable-flies, the parasitic Tachina 
flies, the pomace-flies, fruit-flies, grass-stem flies, brackish-water flies, and 
numerous other kinds not familiar enough to have a vernacular name. To 
get acquainted with some of the more abundant and interesting kinds, and 
to enable us to classify them to subfamiHes (if the whole group is called 
family), we may scrutinize any fly which our key on page 332 leads us to 
call a Muscid, in the hght of the following key: 

(The first posterior cell is the space between the little cross-vein in the middle of 
the wing and the outer margin of the wing. See in Fig. 490.) 

Alulets small Acalyptrate Muscid.i;. 

Alulets large Calyptrate Muscid.e. 

First posterior cell widely open Subfamily Anthomyiin.e. 

First posterior cell narrowly open or closed (Fig. 490). 

Antennal bristle wholly bare Subfamily Tachinin.e. 

Antennal bristle with some distinct hairs. 

Antennal bristle bare near the tip Subfamily Sarcophagin^. 

Antennal bristle plumose or pubescent to the tip. 

Back of abdomen bristly, legs unusually long Subfamily Dexiin.e. 

Back of abdomen not bristly, except sometimes somewhat so near tip. 

Subfamily MusciN^. 

The Acalyptrate Muscidae include a host of small, mostly unfamiliar, 
flies, distributed among a score of subfamilies. We shall refer to a few 

Fig. 483. Fig. 484 

Fig. 483. — House-fly, Musca domestica. (After Howard and Marlatt; three times 

natural size.) 
Fig. 484. — Foot of house-fly, showing claws, pulvilli, and clinging hairs. (Greatly 


of the more interesting kinds in the group after taking up briefly the five 
subfamilies of larger, more noticeable Calyptrate Muscids. 


The Two-winged Flies 

Most abundant, most wide-spread, and most important to us of all the 
Muscid flies are the common house-flies. They belong with some other 
similar forms to the subfamily Muscinas. A number of species may be 
found in houses, but the true house-fly, Musca domestica (Fig. 483), is by 
far the most numerous. Dr. Howard, government entomologist, who has 
paid special attention to the life of house-flies and mosquitoes, because of 
their dangerous disease-germ carrying habits, says that house-flies undoubtedly 
contribute materially in the dissemination of infectious diseases by carrying 
germs in the dirt and filth on their feet, collected during their pilgrimages 
to the contents of cuspidors, slop-pails, and closets. He advocates a definite 
crusade against the house-fly like the one now being undertaken in this 
country against the mosquito. 

Fig. 4S5. 

Fig. 486. 

Fig. 485. — Larva of house-fly, Musca domestica. (After Howard and Marlatt; three 

times natural size.) 
Fig. 486.^Pupa, in puparium, of house-fly, Musca domestica. (After Howard and 

Marlatt; three times natural size.) 

The eggs of the house-fly are laid in horse-manure, occasionally in other 
excrementitious or decaying matter. Each female lays about one hundred eggs. 
These eggs hatch in six or seven hours, and the slender pointed white larvae 
called maggots (Fig. 485) lie in their plentiful food-supply for the five or six days 
necessary for their full growth. They pupate within the last larval skin, which 

thickens and turns brown at the time of pupation 
(Fig. 486). The pupal stage lasts five days, and 
then the fly issues. Its food is liquid and taken 
up by lapping. The "house-fly" that bites is 
not the true house-fly, but usually the fiercely 
piercing stable-fly, Stomoxys calcitrans, another 
member of the subfamily, which looks much like 
Musca and which is a not infrequent visitor in 
the house. 

This stable-fly and another ally of the house- 

moxys calcitrans. (Three fly^ called the hom-fly, are great pests of stock. 

The horn-fly, Hcematobia serrata (Fig. 488), which 

gets its popular name from the habit of clustering, when not feeding, on the 

bases of the horns of cattle, is a European insect that was accidentally brought 

to this country in 1886 or 1887. 

It quickly established itself, and in two years had spread over the eastern 

Fig. 487. — A stable-fly, Stp- 

The Two-winged Flies 


states so widely as o cause much alarm. By 1895 i^ ^^^ spread over all 
of the United States east of the Rocky Mountains. The flies pierce the 
skin and suck the blood, thus causing such an irritation and loss of blood 
that the affected animals cease feeding and soon show great loss in milk or 
weight. The eggs are laid in fresh cow-manure, and the larvcc become full- 
grown and pupate in less than a week. The pupal stage lasts from five 
to ten days. Probably half a dozen generations appear annually. Infested 


-The horn-fly, Hcsmatobia serrata. (After Lugger; natural size indicated 
by Hne.) 

cattle may be smeared with a mixture of resh oil and tar, equal parts, which 
repels the flies, and lime, which kills the larvae, may be thrown on the manure. 
The stable-fly, like the house-fly, lays its eggs in horse-manure, and Dr. 
Howard foresees a curious benefit to result from the gradual increase in the 
use of automobiles in cities, and the corresponding decrease in number of 
horses maintained, in the gradual doing away with the breeding-places of 
house-flies and stable-flies. 

Next to house-flies the commonest ones about houses and outbuildings 
are the bluebottles and blow-flies or flesh-flies. These all lay their eggs 
or deposit living larvae on meat, and, with some other allied species which, 
however, do not all restrict their egg-laying to animal substances, belong 
to the subfamily Sarcophaginas, so named from the flesh-eating habits of the 
larvae or maggots of the best-known species The most abundant flesh- 
fly in this country is named Sarcophaga sarracenicc (Fig. 489), and looks like 
an extra-large house-fly. It gives birth to larvae (hatched from eggs retained 
in the body of the female) which are deposited on fresh meat, sometimes in 
open wounds. The larvae (maggots) feed and grow rapidly, attaining their 
full .size in three or four days. They pupate within the thickened brown last 


The Two-winged Flies 

larval skin, and issue as adults in ten or twelve days after birth. The blow- 
flies and bluebottles, members of this subfamily, have the body steely blue or 
greenish and are great buzzers. The blow-fly, Calliphora erythrocephala, 
has the thorax black and abdomen steely blue. Its eggs are laid on exposed 
meat, fresh or decaying, such egg-infested meat being called ''blown." The 

Fig. 480. — . 

A blow-fly or flesh-fly, SarcopJiaga sarracenicB. 
indicated by line.) 

(After Lugger; natural size 

larvae feed on the juices of the decaying meat and pupate after a few days. 
The pupae enclosed in the thickened brown last larval skin look like 
large smooth shiny brown elliptical seeds, as do indeed the pupa? of all 
Calyptrate ^luscidie. The commonest bluebottle- or greenbottle-fly is Lucilia 

ccBsar, which lays its eggs in 
cow-dung as well as on flesh, 
and which often comes into 
houses, particularly before rain. 
A flesh-fly of serious importance 
is the terrible screw-worm fly, 
Compsomyia macellaria, which 
lays its eggs on flesh, manure, in 
open wounds, and often in the 

Fig. 490. — ^Diagram of wing of Lucilia casar, 
showing venation. 

nasal passages of domestic animals and human beings, entering the nose for 
this purpose while the unfortunate person or animal is asleep. Numerous 
frightful cases of such attacks on persons are recorded, especially from the 
southern states. The lar\-a? fairly eat away the whole inner nose and upper 

The Two-winged Flies 345 

pharynx, causing terrible pain and sometimes death. Indeed, out of twelve 
cases which came to the knowledge of Dr. Richardson, an Iowa physician, 
eleven resulted fatally. As many as three hundred screw-worms were taken 
from the inner nose and region above and behind the soft palate of some 
of the patients. As a pest of domestic animals the greatest injuries have been 
caused in Texas. The eggs are laid in any open wound or in the nose or mouth, 
and the quickly hatching larvae burrow into the adjacent tissues. Cattle and 
hogs are particularly attacked, horses and sheep less often. 

In the states in which sugar-beets are grown some anxiety for the success 
of this new industry — new in this country, that is; sugar has long been made 
from beets in Germany — is felt because of the presence in the beet-fields 
of an obscure little fly, Pegomyia vicina, which maybe called the sugar-beet 
midge. The eggs are laid on the leaves, and in three or four days the tiny 
white larvae hatch and burrow into the soft leaf-tissue. When many of the 
larvae are at work mining the leaves much injury to the plants results. In the 
great sugar-beet fields along the California coast four or five generations 
of this fly appear annually and occasion great loss to the growers. This 
fly belongs to the subfamily Anthomyiinae, to which Muscid group two 
other well-known fly-pests belong, namely, the onion-fly, Phorhia ceparum, 
and the cabbage maggot-fly, Phorbia brassicce. Both these insects in the 
adult stage are small light-gray flies, looking rather like small house-flies. 
The onion-fly lays its eggs on the stems of onion-plants, near the soil, and 
the hatching larvae burrow into the underground bulb, which they soon 
nearly destroy. This fly appears to live on no other plant. The cabbage 
maggot-fly lays its eggs also on the stem just above or even below the ground, 
and the larvae burrow into the roots. Cauliflowers as well as cabbages 
are attacked, and often tens of thousands of acres of these two vegetables 
are destroyed in a single season by this little fly. The best remedy is the 
use of cards cut from tarred paper and bound, collar-like, around the stems 
of the plants. These protecting collars should be put on when the young 
plants are transplanted from the cold frames into the field. Another familiar 
member of this subfamily is the little house-fly, Homalomyia caniailaris , 
smaller, paler, and more conical in shape than the true house-fly. 

Every one who has undertaken to rear butterflies and moths from their 
caterpillars has been compelled to make the acquaintance of certain heavy- 
bodied bristly flies which appear now and then from a cocoon or chrysalid 
in place of the expected moth or butterfly. These are Tachina-flies, and in 
their appearance and parasitic habits are representative of the large sub- 
family of house-fly cousins known as Tachiniinae. The females fasten their 
eggs to the skin of young caterpillars, the hatching larvae burrow into the 
body of their crawhng host and feed on its body-tissues Sometimes the 
caterpillar is killed before it can pupate, but usually not, spinning its cocoon 


The Two-winged Flies 

and pupating with its fatal parasites still feeding inside. But the butterfly 
never issues: in its place buzz out several of these bristly Tachina-flies. 
While their habits arouse our indignation at first acquaintance, and par- 
ticularly if we have set our hearts on rearing a rare moth or butterfly, a 
moment's reflection assures us of the immense good these flies must really 
do. Howard tells of an instance observed by him where the buzzing of 
the swarms of Tachina-flies, hovering over and laying their eggs on the 
hosts of a great army of army-worms, could be heard for a long distance. 

Fig. 491. 

Fig. 492. 

Fig. 491. — A Tachina-fly, Dejeania corpulenta. (One and one-half times natural size.) 
Fig. 492.— Tachinid parasite (at left) of the California flower-beetle, and parasitic fungus, 
Sporotrichum sp. (at right) of same beetle. (Slightly enlarged.) 

He says that a great outbreak of army-worms in northern Alabama in 1881, 
when all crops were threatened with total destruction, was completely frus- 
trated by Tachina-flies. These parasites also attack locusts, leaf-eating 
beetles, and many other injurious insects besides caterpillars, and altogether 
do much to keep in check some of our worst insect-pests. A single species 
of Tachina-fly (Fig. 492) is almost the only check on the destructive flower- 
eating Diabrotica {D. soror) of California, which, if allowed to increase 
unhindered, would soon destroy every blossom in this land of flowers. 

Resembhng somewhat in appearance the Tachina-flies are the so-called 
nimble-flies, constituting the small subfamily De.xiince. Most of the species 
in this country belong to the single genus Dexia and have been little studied. 
The larvae seem to be all parasitic, although the life-history of no species has 
been wholly worked through yet. Beetles and snails seem to be the favorite 
hosts of these flies. 

In the large group of flies, some dingy and obscure in coloration, others 
brightly colored and with beautifully patterned wings, but all small and 
most unfamiliar, called the Acalyptrate Muscidae (that is, the house-fly 
allies with small alulets), we shall not attempt to distinguish the vari- 
ous subfamilies as we have for the Calyptrate Muscids. Dipterologists 

The Two-winged Flies 


recognize some twenty distinct subfamilies (or families, if the group 
Muscidse be looked on as a super-family) of these small flies, but the distinc- 
tions are quite too fine for the general collector to handle. I shall therefore 
simply refer briefly to a few of the more interesting or abundant or economi- 
cally important species in this group. 

Fig. 493. — Red-tailed Tachina-fly, Winthemia 4-pustnluta, a parasite of the army-worm, 
Leucania unipuncta. a, fly, natural size; b, fly, enlarged; c, army-worm, natural 
size, upon which eggs have been laid; d, parasitized army-worms, enlarged. (After 

Of interest because of the extraordinary condition of their eyes are the 
blackish flies called Diopsidae, which have the eyes on conspicuous elon- 
gate lateral processes of the head. These eye-stalks bear also the antennae. 
Only a single species, Sphyracephala brevicornis, has been found in this 
country, and regarding its life-history nothing is known. The flies are to be 
looked for in woodsy places, and particularly on the leaves of skunk-cabbage. 

In the water and cast up in masses along the shores of Mono Lake and 
certain other similar brackish-water lakes in the desert land just east of 
the Sierra Nevada Mountains in CaHfornia may be found, at certain seasons 
of the year, innumerable larvc-e of a small predaceous fly of the genus Ephydra. 
These dead-sea waters support hardly any other animal life, but this fly 
finds the water much to its liking and breeds there with extraordinary fecun- 
dity. The Pai Ute Indians of this region, who, like the flies, have a ques- 
tionable palate, gather these larvae by the bushel, dry them in the sun, and 
use them for food under the name koo-chah-bee. Prof. Brewer of Yale, 
who made a trial of koo-chah-bee, says "it does not taste badly, and if one 


The Two-winged Flies 

were ignorant of its origin it would make a nice soup." Other species of 
Ephydridae occur abundantly in salt-water marshes, the flies living a preda- 

FlG. 494. 

Fig. 495. 

Fig. 494. — Scatophaga sp. (Two and one-half times natural size.) 
Fig. 495. — An aquatic muscid, Tetanocera pictipes, larva, pupa, and adult. 
Needham; two and one-half times natural size.) 


tory life and doing much to reduce the numbers of brackish- water mos- 
quitoes and other small insect-pests. 

One of the great packing-houses of Kansas City, Missouri, once called in 
an entomologist to aid it in fighting a little fly which was causing the packers 
a loss of many thousand dollars annually. This was 
the cheese-skipper fly, Piophila casei (Fig. 496), which 
might almost as well be called the ham- and bacon- 
skipper fly, for the eggs are laid quite as willingly 
on any smoked meat as on cheese. In the packing- 
house swarms of the flies were buzzing about at 
the mouth of the great smoke-shaft from which the 
hams and pieces of bacon were being constantly 
taken to be wrapped and made ready for shipping. 
These flies would dart down and lay their eggs on the 
smoked meat while actually in the wrapper's hands, 
and thus thousands of egg-blown hams and bacon 
sides would be wrapped and sent out. When the cook a thousand miles 
away tears the wrappings from a "piophilized" ham he quickly sends in 
an indignant report to his local meat-supplier, who in turn makes a protest 
to the packer. In time the packer calls for help from an entomologist. 
The larvae of this fly have the odd habit of bending nearly double and 
then with a quick straightening they throw the body some inches into the 
air. Hence the name skipper, commonly applied to it. 

Fig. 496. — The cheese- 
skipper fly, Piophila 
casei. (Five times 
natural size.) 

The Two-winged Flies 


At cider-making and fruit-gathering time, and in vine-growing districts 
at wine-making time, hosts of tiny yellowish-bodied flies, the pomace-flies or 
fermenting fruit-flies, Drosophihdae, may be seen busily lapping up their 
favorite food, the juices of fermenting fruits. 
The most abundant and wide-spread species 
is Drosophila ampelophila, the vine-loving 
pomace-fly. It is a small, clear-winged, 
red-eyed, brownish-yellow, chubby fly 
which lays its eggs on gathered fruits, 
and especially decaying fruit and pomace, 
and also on grapes still hanging on the 
vines if they have been broken somewhat 
by birds. The larva or maggots hatch in 
from three to five days, live in the fruit four days, and lie in the pupal 
stage three to five days, so that a whole life-cycle is gone through in less 

Fig. 497. — Trypeta longipennis. (Two 
and one-half times natural size.) 

Fig. 498 — Larva of cherry-fruit fly, Rhagoletis cingiihta, dorsal and lateral views. 
(After Slingerland; natural size and much enlarged.) 


The Two- winged lilies 

than two weeks. Thus e\'en in the short season of the fruit ripening and 
gathering much injur}- can be and often is done by these little tipplers. 

A much larger group of fruit-flies is the Trypetidas, whose larvae burrow 
in fruits or plant-stems, often producing galls on these latter. The familiar 
spherical swelling or gall on goldenrod stems is the hiding and feeding place 

Fig. 499. — Pupari;i of cherry-fruit fly, Rliugoletis cingiilata. (After Slingerland; natural 

size and much enlarged.) 

of the thick white larvae of Trypeta solidaginis, a pretty fly with banded 
wings. The longer hollow gall which sometimes occurs on goldenrod 
is made by the caterpillar of a small moth, Gelechiq gallcE-solidaginis. 
Some Trypetid species do much injury by burrowing into fruit, as the apple- 
maggot, and the larva of a black-and-white fly with 
banded wings known as Trypeta liidens, whose 
larvK infests Mexican oranges and may sometime 
get a foothold in Cahfornia or Florida. 

Another group of small flies whose larvae are 
responsible for serious injury to growing grain, 
meadows, and pasture grasses are the Oscinidae, 
or grass-stem flies. The adults are commonly taken 
by collectors when beating or sweeping in meadows 
Fig. 500. — An aquatic and pastures. The flies are minute but plump, 

muscid, Sepedon fasa- ^^^^ ^j.^ variously colored, sometimes blackish, 

pennis, larva, pupa, and . . 

adult. (After Needham ; Sometimes yellowish. They are so small that they 

two and one-half times often get into one's eyes in their swarming-time, 

and are said to cause a prevalent disease of the 

eyes in the South. The thick cylindrical little larvae of several species of 

Oscinis hve in the stems of wheat, barley, oats, rye, and grass. The larva 

of Chlorops similis burrows in the leaves of sugar-beets, and another 

The Two-winged Flies 351 

species of the genus is the notorious "frit-fly," one of the chief grain- 
pests of Europe. 


Bird-collectors occasionally find on their sj)ecimens curious flat-bodied 
insects with leathery skin and a single pair of wings, which are obviously 
parasites on the body of the birds. Owls and swallows seem especially 
infested. Similar parasitic insects, but wingless, are also found on sheep, and 
a winged form is not uncommon on horses. These degraded insects are 
flies of the suborder Pupipara which are commonly known as bird-ticks, 
sheep- and horse-ticks, etc. The animals more rightly entitled to the name 
"ticks" are really not true insects, but belong with the scorpions, spiders, 
and mites in the class Arachnida. They have four pairs of legs and are always 
wingless. Such true ticks are the leathery-skinned cattle-ticks, dog-ticks, 
and wood-ticks. 

The degraded Diptera belonging to the suborder Pupipara, and also 
called ticks, have of course three pairs of legs and some are winged. Their 
name Pupipara comes from the curious circumstances of their birth. The 
female does not deposit eggs outside her body, but gives birth to young which 
are just ready to assume the pupal stage at the time of their appearance. 
In the case of one species, the sheep-tick (Melophagus), whose development 
has been carefully studied, the female has four egg-tubes each of which 
produces a single germ-cell at a time. Of these four egg-cells three remain 
small, while one becomes large and develops into an embryo. This embryo 
lies in the unpaired wide vagina of the female, soon casts ofT its egg-envelopes, 
and is nourished as a growing larva by a secretion from two pairs of glands 
opening into the vagina of the mother. Here the headless, footless larva 
lies and grows until it is about ^ inch long, when it is born and immediately 
pupates. The development of the other Pupipara, as far as studied, is 
similar to that of the sheep-tick. 

The suborder includes three famihes, as follows: 

With compound eyes; sometimes with wings. 

(Bird-, sheep-, and horse-ticks.) Hippoboscid^. 
Without compound eyes, always wingless. 

Halteres present; on bats (Bat-ticks.) Nycteribiid^. 

Halteres absent; on honey-bees (Bee-lice.) Braulid^. 

Of the Hipposcida; the sheep-tick, Melophagus ovinus, already referred 
to, is common and familiarly known. It is wingless, and can crawl readily 
about through the wool next to the skin. With its strong proboscis, com- 
posed of two hard pointed flaps, it punctures the skin and sucks blood from 
its host's body. The horse-tick, Hippobosca equina (Fig. 501), is winged. 
There are several species of this family found on birds. Oljersia americana 


The Two-winged Flies 

is a yellowish winged species common on owls, some hawks, and the ruffed 
grouse. Swallows are often infested, and I have taken bird-ticks from half 
a dozen other kinds of birds. A careful search for these curious insects 
will certainly make known numerous new species. 

(.-Xfter Lugger; natural 

Fig. 501. — A horse-tick or forest-fly, Hippohosca equina. 

length J to J inch.) 

The genus Lipoptena includes a few known species found on mammals 
which are winged for awhile, but later cast or bite off the wings. They 
probably fly about in their search for a host, after finding which they remove 
their wings and remain for the rest of their life on this host individual. Lip- 
optena cervi is a species found on deer. 

Fig. 504. 

Fig. 502. Fig. 503. pic. -03. Bat-tick, Nycterihia sp. 

Fig. 502. — Sheep-tick, Melophagus ovinus. Nat. size \ in. 

Fig. 504. — A bee-louse, Braula sp. (After Sharp; much enlarged.) 

The bat-ticks, Nycteribiidse (Fig. 503) , are curious long-legged, wingless, 
small spider-like creatures about \ inch long or less, which look as if the 

The Two-winged Flies 353 

upper were the under surface. The head is narrow and Hes back on the 
dorsum of the thorax, and the pro thorax rises from the upper instead of 
anterior aspect of the mesothorax. They are found only on bats and are not 

The strange minute insect, jV inch long, found clinging to the thorax 
of queen and drone honey-bees and known as the "bee-louse," Braula 
ca'ca (Fig. 504), is the only species known of the family Braulidae. Its legs 
are rather short and stout, and each ends in a pair of comb-like brushes. 


The fleas are blood-sucking parasites of mammals and birds which were 
long classified as a family (Pulicidae) of the Diptera, being looked on as 
wingless and otherwise degenerate flies. But they are now given by ento- 
mologists the rank of an order, called Siphonaptera, subdivided into three 
famihes of its own. Nearly one hundred and fifty species of fleas are known 
in the world, of which about fifty are recorded from this country. They have 
been taken from the domestic dog, cat, rat, and fowls, and from various wild 
animals, such as several rabbit and squirrel species, the lynx, weasel, mole, 
mountain-rat, shrews and mice, prairie-dog, woodchuck, opossum, etc. 
Rothschild has recently described a new flea species from the grizzly bear 
(British Columbia). But from the great majority of our wild mammals fleas 
have not yet been recorded, although undoubtedly most of them are infested. 
Baker, who has recently published a monograph* of the known North 
American species, suggests that particularly interesting forms will probably 
be found on bats. One flea species, Pulex avium, has been taken from several 
kinds of birds, and two or three other fleas are recorded from bird hosts. 

The 'pecuhar structural characteristics of fleas are their winglessness, 
the extraordinary lateral compression of the body, and the curious modifica- 
tion of their mouth-parts for effective piercing and blood-sucking. The an- 
tenna lie in little half-covered grooves, extending down and back behind 
the eyes; they can be lifted or stretched up whenever needed. Each antenna 
is composed of three segments, the terminal one, however, being spirally or 
transversely lined or grooved and variously shaped, so that it appears to be 
composed of several segments. The mouth-parts consist of a pair of needle- 
like mandibles, a pair of slender grooved labial processes, probably the 
palpi, a pair of short, broad, flattened maxillae, each with a short antenna- 
like palpus at its tip, and an unpaired needle-like hypopharynx. The needle- 
like parts serve for piercing and the grooved labial processes for sucking. 
Regularly arranged over the body are (in most fleas) many series of stiff, 
spine-like hairs, often unusually conspicuous and strong on the head and 

* Baker, C. F. A Revision of American Siphonaptera. Proc. U. S. Nat. Mus., vol. 
xxvii, 1904, pp. 365-469. 


The Two-winged Flies 

thorax. The head is ridiculously small and malformed, so that a flea under 
the microscope always suggests an idiotic (microcephalous) creature. But 
if its insidious attack and brilliant tactics in retreat be due to wit, this 

Fig. 505. — Dog- and cat-flea, Ctenocephalus canis. (After Lugger; much enlargeci.) 

small-headedness is truly deceptive. However, our modern mechanical 
theories of reflex action, negative phototropism (repulsion by light), etc., 

Fig. 506. — The house-flea, Pw/ex ^Vritew5. ^, larva; 5, pupa; C, adult. 
(After Beneden; much enlarged.) 

allow r.s to give the elusive flea little credit for its ingenuity; we must look 
on it ..s an unusually well-made and smoothly-working organic machine. 

The Two-winged Flies 355 

While the adult fleas are commonly seen, particularly in lands of soft 
climate, like Italy and California, in immature form these insects are wholly 
unfamiliar. The larva) (Fig. 506) are small, slender, white, footless, worm- 
like grubs, with the body composed of thirteen segments, the first being the 
small brown head bearing short antenna) and biting mouth-parts, but no 
eyes. The larvce seem to live on dry vegetable dust, the excreta of adult 
fleas, and other organic detritus. The larval life varies much in duration 
in different species, and even in the same species under varying conditions. 
In our commonest species, the cat- and dog-flea, Pergande has found the 
larval hfe to last only one or two weeks, the whole development from egg to 
adult being completed sometimes in a fortnight. When full-grown the 
larva spins (usually) a thin silken cocoon in the dust or htter in which it hes, 
within which it pupates. 

The parasitic habits of fleas vary from a very temporary character to one 
approaching permanence. In such forms as the human flea and the dog- 
flea no stage of the immature life is passed on the body of the host (although 
the eggs of the dog-flea are usually laid on the hairs of the host, but so loosely 
attached that they fall off before the larvae emerge), but in the burrowing 
kinds hke the "chigoe" or "jigger," where the females become completely 
encysted in the skin of the host, the young hatch in the tumor, and unless 
carried out by pus probably develop there. But taken altogether the fleas 
are to be considered as belonging to the category of "temporary external 


The species known in this country represent two families which may be 
separated by the following key: 

Small fleas with proportionally large head ; female a stationary parasite with worm- 
like or spherical abdomen, burrowing into flesh of the host; labial palpi 
i-segmented; no "combs" of spines on head, thorax, or abdomen. 


Larger fleas with proportionally small head; adults active temporary parasites, 
with abdomen always compressed; labial palpi 3- to 5-segmented; head, 
thorax, or abdomen often with "combs" of spines Pulicid.^. 

Of the SarcopsyUida) but two genera are known, one, Sarcopsylla, includ- 
ing the common jigger-flea, infesting various mammals and man in the 
tropics and probably occurring in Florida and southern Texas, and Xes- 
topsylla, the common chicken-flea, being distinguished by having the head 
not angularly produced. 

The jigger-flea, or chigoe, Sarcopsylla penetrans (not to be confused with 
a minute red mite, common on lawns, which burrows into the skin and is 
also called "jigger" or "chigger"), was described by Linnaeus in 1767 and 
has been commonly known as a pest of man in tropical and sub-tropical 
countries ever since. It also infests many domestic animals, as the dog, cat, 

35^ The Two-winged Flies 

horse, cow, sheep, etc., as well as birds. The male jigger-fleas hop on or 
off the host as other fleas do, but the females, when ready to lay eggs, burrow 
into the skin, especially that of the feet, and produce a swelhng and later 
a distinct ulcer, sometimes so serious as to result fatally. The remedy is 
(as also for the chigger-mite) the pricking out entire, with a needle or knife- 
point, of the pest as soon as its presence is detected. The bursting of the 
body of the female in the skin, with the release of its eggs, is likely to result 
seriously. When domestic animals are attacked it is difficult to fight the 
pest. The liberal use of pyrethrum on the rubbish or dust in which the 
young stages are developing is recommended. The hen-flea, XestopsyUa gal- 
linacea, first described from Ceylon, sometimes becomes a serious pest of 
fowls in warm regions. The females of the hen-flea burrow into the skin of 
the fowl and lay their eggs in the small tumor which forms about them. 
This pest has been known in the Southern United States since about i8go 
and is a common pest from Florida to Texas. 

The second family, Pulicidae, includes all the other fleas, none of which 
burrows into the skin. The various species range in size from ^^ inch (Anomi- 
opsylhis niidatus, found on a mouse in Arizona) to \ inch {Ceratophyllus 
stylos Its, taken from Haplodon in Oregon), but all fairly similar in shape 
and appearance to the familiar house-fleas. They are grouped in nine 
genera, of which Pulex is much the largest and includes the human flea 
and the cat- and dog-flea, the two species to which the house-infesting pests 
belong. The human flea, Piilex irritans, was described by Linnaeus in 
1746. It is known all over the world, and often becomes a serious pest. 
In this country it is probably not so commonly met with in houses as the 
cat- and dog-flea, Ctenocephalns canis, from which it may be readily dis- 
tinguished by its lack of combs of spines on the back of the head and 
prothorax. The eggs of irritans "are deposited in out-of-the-way places, 
in the dust or lint under carpets, and the larvae are said to feed upon the 
particles of organic matter which may be found in such localities." Raillet 
states that each female deposits eight to twelve eggs from which larvae hatch, 
in summer, in from four to six days, become pupae eleven days later, and 
after about twelve days in this stage become adult. In winter, in warmed 
houses, the whole development takes about six weeks. The cat- and dog- 
flea lays its eggs on or among the hairs of an infested animal, but the 
eggs drop to the floor or ground as the animal moves about, and the larvae 
live in the dust, feeding on whatever bits of organic substance they can find 
there. Larvae placed on dust with birds' feathers mixed with dried blood 
developed perfectly. Others put on the sweepings of a room developed 
as well. These fleas are especially abundant and troublesome in houses 
in the East in damp summers. As flea-larvae will not develop successfully 
in places where they are often disturbed, much sweeping and scrubbing 

The Two- winged Flies 357 

will keep them down. Mats and places where dogs and cats He down should 
be kept well dusted with pyrethrum. (Buhach is the trade name for this 
insecticide, which is not injurious to man or domestic animals.) Where 
fleas get a foothold in a neglected room or cellar, the remedy used by Profes- 
sor Gage in the basement of one of Cornell University's buildings might be 
tried; i.e., tying sheets of sticky fly-paper, sticky side out, around the legs 
from foot to knee of the janitor or a cheap boy and having him tramp for 
several hours around in the room! 

Of the various other flea species, the only ones that come into special 
relation with man are the rat-fleas. The proof that rats are active agents 
in the dissemination of the dreadful bubonic plague, and the behef of some 
pathologists that the disease-germs may be transmitted from rats to man 
by the bites or punctures of rat-fleas, gives this insect a special interest like 
that attaching to the malaria- and yellow-fever-dissminating mosquito and 
the germ-carrying house-fly. Baker pertinently calls attention to the fact 
that the rat-fleas of this country are only remotely related to Pulex irritans 
and Ctenocephalus canis, the two species that bite human beings, while the 
fleas that infest rats in the tropics are, on the contrary, very nearly related to 
the man-infesting kinds. The prevalence of the bubonic plague in tropical 
countries and its rarity with us may be connected with this difference in the 
rat-flea kinds. 



OTHS and butterflies are the insects most 
favored of collectors and nature lovers; a 
German amateur would call them the "Lieb- 
lings-insekten." The beautiful color patterns, 
the graceful flight and dainty flower-haunting habits, and the interesting 
metamorphosis in their life-history make them very attractive, while the com- 
parative ease with which the various species may be determined, and the 
large number of popular as well as more technical accounts of their life 
which are accessible for information, render the moths and butterflies most 
available, among all the insects, for systematic collecting and study by 

Despite the large number of species in the order (6622 are recorded in 
the latest catalogue of the North American forms) and the great variety in 
size and pattern, the order is an unusually homogeneous one, even a begin- 
ning student rarely mistaking a moth for an insect of any other order, or 
classifying a non-lepidopterous insect in this order. A few aberrant species 
are wingless (females only) and a few (certain "clear-winged" species) have 
a superficial likeness to wasps and bumblebees, but the general habitus of 
any Lepidopteron, let alone the readily determinable and absolutely diag- 
nostic character of the scale-covering on the wings, usually indicates unmis- 
takably the afluiities of any moth or butterfly. 

The diagnostic structural characters are the (already mentioned) pres- 
ence on upper and lower sides of both wings (as well as over the surface of 
the body) of a covering of small sy-mmetrically formed scales, which are 
modified hairs, and to which all of the color and pattern of the insects are 
due. In Chapter XVII will be found a detailed account of these scales, 
e.xplaining their structure, their origin, and how they produce the color pat- 
terns. The wings themselves are almost always present (in two pairs), the 
fore wings larger than the hind wings, and with a characteristic venation, 
in which the modifications, though small, are yet so constant and definite 
that they are used successfully as the principal basis for the classification 
of the order into famihes. Another characteristic is the highly modified 
and peculiar condition of the mouth-parts. While in some species the mouth- 
parts are rudimentary (atrophied) and evidently not functional, in most 
there is a well-developed slender flexible sucking proboscis (Fig. 509) com- 




i = Junonia coenia. 
2=Iphidides ajax. 
3=Epargyieus tilyrus. 
4= Cyan iris pseudargiolus. 
5 = Ancy loxypha n umitor. 
6=Papilio turnus. 
7 = Nathalie iole. 
8=Parnassiu3 smintheus. 
9=Tlieda halesus. 
io= Zerene ctesonia 


Mary IVellman, del. 

The Moths and Buttertiies 


posed of the two greatly elongate maxillae, so apposed that a groove on the 
inner face of one fits against a similar groove on the inner face of the other, 
the two thus forming a perfect tube (Fig. 510). This sucking proboscis, when 
extended, may protrude five or six 
inches, as in some of the sphinx- 
moths, or only a fraction of an inch, 
as in the small moth "millers," but 
when not in use it is so compactly 
coiled up, watchspring-like, under 
the head, and so concealed by a 
pair of hairy little tippets (the labial 
palpi) which project up on each 
side of it that it is nearly invisible. 
Of the other mouth - parts, the 
upper lip (labrum) and under hp 
(labium) are greatly reduced and 
are not movable and flap-like as in 
most insects, while the mandibles 
are either wholly wanting or, as in 
the sphinx-moths and some others, 
represented only by small immov- 
able functionless rudiments. The 
palpi of the maxillae are also either 
wholly wanting or present as mere 
rudiments. The foregoing descrip- 
tion of the mouth-part conditions 
is true for the great majority of 
Lepidoptera, but among the lowest 
(oldest or most generalized) moths 
some interesting examples of much 
less specialized conditions occur. 
Indeed in one family of minute 
moths, the Eriocephalidas, all the 
usual parts of a typical insect 
mouth are present and in a condi- 
tion fitted for biting and chewing 
and in all ways wholly comparable 
with the condition in such biting 
mandibles are movable 

Fig. 507. — A trio of apple tent -caterpillars, 
larvas of the moth Clisiocampa americana. 
These caterpillars make the large unsightly 
webs or tents in apple-trees, a colony of 
the caterpillars living in each tent. (Photo- 
graph from life by Slingerland; natural size.) 

insects as the locusts and beetles; the 
and truly jaw-like, the maxillas short and rlso 
jaw-like and provided with several-segmented palpi, while both labrum 
and labium are truly lip- or flap-like and fully movable, the labium 
bearing 3-segmented palpi. Between this most generalized condition 


The Moths and Butterflies 


Fig. 508. — Bit of wing of monarch but- 
terfly, Anosia plexippiis, showing scales; 
some scales removed to show the inser- 
tion-pits and their regular arrangement. 
(Greatly magnified.) 

stage. The immature staees of 

and the extreme speciaUzation of the butterfly's mouth an interesting and 
illuminating gradatory series is dis- 
coverable by examining moths of suc- 
cessively more specialized character. 
The development of moths and 
butterflies shows the usual character- 
istics of devel- 
opment with 
complete meta- 
morphosis, the 
larval or cater- 
pillar stage be- 
ing quite dis- 
similar from 
the pupal or 
chrysalid stage, 
and that in 
turn from the 

adult or imaginal stage. 'I'he immature stages 
Lepidoptera are more familiar than those of any other 
order; we have all seen, and recognized for what they 
are, the caterpillars and chrysalids of various moths 
and butterflies. The great silken cocoons found on 
orchard-trees in winter-time are known to contain 
the pupae of giant moths, as the 
Cecropia, the Polyphemus, and others, 
while the soft-bodied green tomato- 
worms are as well known to be the 
young (larvae) of the hawk-moths. 
As a matter of fact the young stages 
of no other of the insects with com- 
plete metamorphosis are so nearly 
unmistakably characterized by their 
common possession of certain well- 
defined features. The larvae or cater- 
pillars, for example, with very few 
exceptions, possess, in addition to 
three pairs of jointed legs on the first 
three segments behind the head, 
from three to five pairs of short 
fleshy unjointed legs or feet called 
prop-legs, on certain abdominal seg- 

FiG. 509. — Sucking-proboscis of a sphinx- 
moth; at left the proboscis is shown 
coiled up on the under side of the head, 
the normal position when not in use. 
(Small figure, natural size; large figure, 
one-half natural size.) 

The Moths and Butterflies 



ments; one of these pairs is on the last segment and four, which is the num- 
ber present in all except the inchworms or loopers (larvae of the Geometric! 
moths), are on the sixth, seventh, eighth, and ninth segments behind the 
head. The inchworms have prop-legs only (with a few exceptions) on the 
ninth and last segments. These 
prop-legs, together with the striped 
or hairy body- surface, make a 
moth or butterfly larva almost as 
readily recognizable for what it is 
as the scale-covered wings make 

Fig. 510. Fig. 511. Fig. 512. 

Fig. 510. — Cross-section of sucking-proboscis of milkweed-butterfly, Anosia plexippus; 

see tubular cavity, c, formed by apposition of the two maxillae, tr., trachea; n., nerve; 

m., muscles. (After Burgess; greatly magnified.) 
Fig. 511. — Bit of maxillary proboscis of milkweed-butterfly, Anosia plexippus, showing 

arrangement of muscles in the interior; these muscles serve to coil up or to extend 

the proboscis; see groove on inner face of maxilla. in., muscles; tr., trachea; 

71., nerve; c, groove. 
Fig. 512. — Diagram of arrangement of pharynx, oesophagus, etc., in interior of head 

of monarch butterfly, Anosia plexippus, showing means of producing suction in 

the proboscis, oe., oesophagus; t?»?., dorsal muscle; /.w., frontal muscle; c/., clypeus; 

hyp., hypopharynx; s.d., salivary duct; ep., epipharynx; mx., maxilla. 

the adult moth or butterfly distinguishable from any other kind of insect. 
The chrysalids with their hard shell, but with the folded antenuce, legs, and 
wings of the enclosed developing adult always indicated, are also hardly to 
be mistaken for the pupae of any other orders, while even the eggs, when ex- 
amined under a magnifier, mostly reveal their lepidopterous parentage by 
the beautiful fine sculpturing of the shell (Fig. 67). As will be noted 
from a perusal of the accounts of the life-history of various familiar and 
representative moths and butterflies given in the following pages, there is 
much variety in the means shown of protecting the defenceless pupae; some 
are subterranean, the leaf-feeding larvae crawling down from tree-top or 
weed-stem and burrowing into the ground before pupation; others are 
enclosed in a tough silken cocoon spun by the larva before making its 
last moult; while those which are not protected in one or the other of these 
ways either lie in concealed spots under stones or in cracks of the bark, 
etc., or are so colored and patterned that they blend indistinguishably with 
the object against which they are suspended. The larvae have also their 


The Moths and Butterflies 

various means of defence; the hairy ones are an uncomfortable mouthful 
for a bird, the naked and brighdy marked ones usually contain an acrid 
and distasteful body fluid, while still others find protection in a color pattern 
harmonizing with their habitual environment. 

The food-habits of the larvae make of many of them serious pests of 
our growing crops. Most are leaf-eaters and all are voracious feeders, so 
that an abundance of cutworms or army- worms or maple-worms or tomato- 
worms always means hard times for their favorite food-plants, which are 
too often growing grain and 
vegetables, and leafing or- 
chard and foliage trees. f ( /f ) niAii)])^-"'"'^'^' 
Others attack fruits, as that ( ^^f\"^^' / y/%r^'^^ 
dire apple pest, the codlin- 
moth larva ; while still others '" p^- '""''^ /ff\y\f)\' ^P- 

Fig. 513. 

Fig. 514. 

Fig. 513. — Front of head, with scales removed, of sphinx-moth, showing frontal sclerites 
and mouth-parts, f/^., epicranium; 5!(., suture; c/., clypeus; ^e., gena or cheek; />/., 
piUfer of labrum; md., mandible. Between the two pilifers the base of the sucking- 
proboscis composed of the apposed maxillae is seen. (Much enlarged.) 

Fig. 514. — -Diagram showing mouth-parts of Lepidoptera. Figure in upper left-hand 
corner, head, with scales removed, of Catocala sp.: cl., clypeus; ge., gena or cheek; 
mx.p., maxillary palpus; pj., pilifer of labrum. In upper right-hand corner, ventral 
aspect of head of Catocala sp.: mx.p., maxillary palpus; ge., gena or cheek; mx.b., 
base of maxilla; gu., gula; Im., labium; Ip., basal segment of labial palpus. In 
lower left-hand corner, frontal aspect of head, with scales removed, of sphinx- 
moth, Protoparce Carolina: ep., epicranium; cl., clypeus; lb., labrum; pj., pilifer 
of labrum; md., mandible; ge., gena or cheek. In lower right-hand corner, ffont 
of head, with scales removed, of monarch butterfly, Anosia plexippus lb., labrum; 
g., gena or cheek; pj., piUfer of labrum. (Much enlarged.) 

are content with dry organic substances, as the larvae of clothes-moths, meal- 
moths, and the like. For all of this kind of feeding very different mouth- 
parts are needed from the delicate sucking-proboscis characteristic of the 
adults, and the lepidopterous larvae are all provided with well-formed jaw-like 
mandibles and other parts going to make up a biting mouth structure. The 
larval eyes are simple ones, not compound as in the adults; the antennae 
are short and inconspicuous, not large and feathered as in the moths, or 
long and thread-like, with knobbed tip, as in the butterflies. Altogether the 

The Moths and Butterflies 


lepidopterous larva is a well-contrived animal for its especial kind of life, 
which is as different as may be, almost, from that which it will lead after 
it has completed its metamorphosis. Always when one reads or hears of 
injurious moths or butterflies it should be kept clearly in mind that the 
injuries, to crops or fruit or woolen clothing or what not, are caused by the 
moth or butterfly in its larval 

stage and never by the flut- 
tering nectar-sipping adult. 
The sole compensation, 
other than the rather imma- 
terial though perhaps not 
less real one afforded us 
through our jesthetic ap- 
preciation of the beauty 
and attractive, apparently 
care-free, flitting about of 


Fig. 517. 

Fig. 518. 



515. — Front of head of larva of tussock-moth, Notolophus leucostigma. ant., antenna; 

md., mandible; vjx., ma.xilla; mx.p., maxillary palpus; li., labium. (Much enlarged.) 
516. — Front of head of old larva of tussock-moth, Notolophus leucostigma, with 

head-wall dissected away on right-hand side to show forming adult mouth-parts 

underneath, l.ant., larval antenna; ant., adult antenna; , larval mandible;, larval ma.xilla; i.nix., adult maxilla; lb., larval labrum; /./i, larval labium. 

(Much enlarged.) 
Fig. 517- — Developing adult head dissected out from head of larva of tussock-moth, 

Notolophus leucostigma. ant., antenna; mx., maxilla; li.p., labial palpus. (Much 

Fig. 51S. — Head of tussock-moth, Notolopliw; leucostigma; showing adult antenna; and 

mouth-parts, mx., maxilla; li.p., labial palpus. Note that the two maxilla; are 

not locked together to form a sucking-proboscis, the mouth-parts of this moth being 

rudimentary and not capable of taking food. (Much enlarged.) 

the butterfly, which the Lepidoptera make for their often disastrous toll on 
our green things, is the prodigal gift of silk made by the moth species known 
as the mulberry or Chinese silkworm. Thoroughly domesticated (the wild 
silkworm species is now not even known), this industrious spinner produces 
each year over one hundred million of dollars' worth of fine silken thread 


The Moths and Butterflies 

ready for the loom. In Italy and Japan nearly every country household has 
its silk-rooms in which thousands of the white "worms" are carefully fed and 
tended by the women and children, and from which comes enough raw silk 
to furnish a good share of the annual income of each of these households. 

The reader who would undertake the collecting of moths and butterflies, 
or the rearing of caterpillars in home " crawleries," is referred for some 
specific directions for this work to the appendix of this book, p. 635 et seq. 

The order Lepidoptera may be most conveniently divided into two prin- 
cipal subgroups (suborders they are often called), namely, the Heterocera, 

Fig. 519. — Larva of obsolete-banded strawberry leaf-roller, Cacoecia obsoletana. (Photo- 
graph from life by SUngerland; natural size in lower corner and twice natural size 

or moths, and the Rhopalocera, or butterflies. All butterflies have antennae 

which are slender (filiform) for most of their length, but have the tip expanded 

or thickened, forming an elongate spindle-shaped dilation or "club"; the 

moths have their antennae variously formed, as wholly filiform, pectinate, 

The Moths and Butterflies 


Fig. 520. — Pupa of obsolete-banded strawberry leaf-roller, Cacoccia ohsoletana. (Photo- 
graph from life by Slingerland; natural size a little more than one-half inch.) 

Fig. 521. — Moths of the obsolete-banded strawberry leaf-roller, Cacoecia obsoletaiia, 
male above, female below. (Photogt^aph from life by Slingerland; natural size.) 


The Moths and Butterflies 

The Moths and Butterflies 


etc., but never showing the characteristic swollen-tipped or clubbed con- 
dition of the butterflies. The moths, too, are mostly night or twilight flyers, 
while the butterflies go abroad in sunlight only. Scientific students of Lepi- 
doptera do not give the butterflies a classific value equivalent to that of the 
moths taken altogether, but rather rank them as a group more nearly equiva- 
lent to a single superfamily of moths, as, for example, the superfamily Satur- 
niina, which includes all our great silkworm-moths, Cecropia, Luna, Prome- 
thea, Polyphemus, etc., etc. However, the more familiar and readily made 
subdivision of the order into moths and butterflies is more convenient and 

Fig. 523. — ISIoth and cocoon cut open to show pupa of Samia cecropia. (After Lugger; 

slightly reduced.) 

quite as informing for our purpose, so we shall adopt it, taking up the moths 
first, as including the more generalized members of the order. There are many 
more moth than butterfly families, the numbers represented in this country being 
44 to 5. By reference to the following key adapted from Comstock aln ost any 
North American moth can be traced to its proper family. 

This key does not include a few of the smaller families whose members are very few 
and are rarely taken by collectors. Some of these moths are, however, referred to in 

368 The Moths and Butterflies 

the systematic account of the families which follows later. To use the key requires 
an acquaintanceship with the plan of venation in the wings and the nomenclature of 
the veins. This may be got from an inspection of Fig. 525, and by referring to the 
various other figures illustrating the typical venation for the various important families. 
To see clearly the veins, a necessary prerequisite to using the key, a few drops of ether 
should be put on the outstretched wing of a spread specimen and this held so that bright 
light, as from a window or lamp, may pass through the wing to the eye. For a few 
moments (until the evaporation of the ether) the covering-scales will be transparent 
and the number and course of the veins plainly visible. The ether will not injure the 
specimen at all. If duplicate specimens are available, the fore and hind wings of one 
side may be removed and placed in a watch-glass or small saucer containing Eau de 
Labarraque (to be obtained of a druggist), when the scales will be bleached perfectly 
transparent. The wings may be then washed and mounted on glass sHdes with glycerine 
jelly and thus be made available for inspection at any time. 

A. Moths which have a thin lobe-Hke process (jugum) projecting backward from the 
base of the fore wing, which holds fore and hind wings together when they are 
outstretched; veins similar in number and arrangement in both wings (Fig. 526). 

(The Jugatse.) 
B. Very small moths, not more than one-fifth inch long. 

MiCROPTERYGiD^ and Eriocephalid.e. 

BB. Moths from one-half to one inch long (The Swifts.) Hepialid^. 

AA. Moths whose wings are not united by a jugum but by a frenulum (Fig. 533), and 
in which the veins in the hind wing are less in number than in the fore wing. 

(The Frenatse.) 
B. Hind wings with fringe on hinder margin as long as the width of the wing; 

hind wings often lanceolate in shape Superfamily Tineina (part). 

BB. Hind wings with narrow or no fringe, and not lanceolate in shape. 

C. Wings fissured, i.e., divided longitudinally into several narrow parts. 
(Plume-moths.) Pterophorid^ and ORXEODiD.i;. 
CC. Wings not fissured. 

D. Fore wings very narrow; part of the hind wings always, and of 
the fore wings often, clear, i.e., without scales. 

(Clear-winged moths.) Sesiid.^. 
DD. Wings all covered with scales or, if partly clear, the fore wings broad. 
E. Hind wings with three anal veins. 

F. Subcosta and radius of hind wings close together or fused 
beyond the discal cell (Fig. 533). 

Superfamily Pyralidina. 
FF. Subcosta and radius of hind wings widely apart beyond 
the discal cell. 

G. Small; palpi usually prominently projecting; fringe on 
inner angle of hind wings longer than on rest of margin. 
H. Second anal vein of hind wings forked at the 

base (Fig. 539) Superfamily Tortricina. 

HH. Second anal vein of hind wings not forked 

at base Superfamily Tineina (part). 

GG. Medium or large; palpi not conspicuously project- 
ing beyond the head and fringe on inner angle of 
hind wings only slightly or not at all longer than 
on rest of margin. 

The Moths and Butterflies 369 

H. Subcosta and radius of hind wings fused nearly 
to end of the discal cell (Fig. 553). 

I. Small black moths. 

(Smoky-moths.) Pyromorphid^ (part). 

II. With long, curling, light-colored or brown 
woolly hairs 

(Flannel-moths.) Megalopygid.e. 
HH. Subcosta and radius of hind wings distinct 
or only slightly fused. 

I. Anal veins of fore wings anastomosing so 
as to appear as a branched vein (Fig. 552). 

(Bag-worm moths.) Psychid.^. 

II. Anal veins not anastomosing. 

J. Vein m^ of fore wings arising from the 
discal cell nearly midway between 
veins Wj and m^ (Fig. 603). 

(Silkworm-moths.) Bombycid^. 

JJ. Vein m^ of fore wings rising from discal 

cell nearer to cubitus than to radius, 

so that cubitus appears four-branched 

(Fig. 548). 

(Carpenter-moths.) CossiD^. 
EE. Hind wings with less than three anal veins. 

F. Fore wings with two distinct anal veins or with these two 
veins partly fused so as to appear like a single branched vein. 
G. The two anal veins distinct (Fig. 553). 

Pyromorphid.e (part). 

GG. The two anal veins partly fused and appearing like 

a single branched vein (Fig. 552). PsYCHiDiE (part). 

FF. Fore wings with but one complete anal vein (rudiments of 

one or two others sometimes present). 

G. Frenulum present. 

H. Hind wings with subcosta and radius 
apparently distinct, but connected by a strong 
oblique cross-vein; moths mostly with narrow, 
long, strong front wings and small hind vwngs. 
(Sphinx- or hawk-moths.) Sphingid^. 
HH. Hind ^vings with subcosta and radius either 
distinct or fused, but not connected by an 
oblique cross-vein. 

I. Vein m..^ of fore wings closer to radius than 
cubitus, cubitus being apparently three- 

J. Subcosta of hind wings extending 
from base to apex of wing in a regular 
curve (Fig. 560); moths with heavy 
abdomen and rather narrow strong 
fore wings. 

(The prominents.) Notodontid.^. 

JJ. Subcosta of hind wings with its basal 

part making a prominent bend into the 

370 The Moths and Butterflies 

humeral angle of the wing (Fig. 567); 

moths mostly with slender abdomen 

and rather broad dehcate fore wings. 

Superfamily Geometrina. 

II. Vein Wj of fore wings more closely joined 

to cubitus than to radius, so that cubitus 

is apparently four-branched. 

J. Subcosta of hind wings distinct from 

radius, or the two fused for a very 

short distance near the base of the 

wing (Fig. 584). 

K. Day-flying moths that are black 
with large white or yellow patches 
on the wings, or with white front 
wings margined with brown, and 
having the hind wings pale yellow. 
(Wood-nymph moths.) Agaristid^ and Pericopid.e. 
KK. Not such moths. 

L. Ocelli absent; antennae pec- 
(Tussock-moths.) Lymantriid^. 
LL. Ocelli present or, if absent, 
with simple antennae. 
(Owlet-moths.) Noctuid.e. 
JJ. Subcosta of the hind wings fused with 
radius for one-fifth or more of the 
length of the discal cell. 
K. Subcosta and radius of hind wings 
fused entirely or vnth only the tips 
separate (Fig. 591). . .Zyg^nid.e 
KK. Subcosta and radius of hind wings 
united for about one-half their 
length, of more, but usually 
separating before the apex of the 
discal cell (Fig. 597). 
L. Ocelli present. 

(Tiger-moths.) Arctiid^e. 

LL. OceUi absent. 

(Footman-moths.) Lithosiid.e. 

GGo Frenulum absent; the humeral angle of the hind 

wings largely expanded and serving as a substitute 

for the frenulum (Fig. 600). 

H. Cubitus of both wings apparently four-branched 
(Fig. 600). (Tent-caterpillar moths et al.) 

HH. Cubitus of both wings apparently three- 
branched; robust moths with broad wings (Fig. 
603). (Giant silkworm-moths.) Saturniina. 

The jugate moths include but two families, the Micropterygida^ and 
Hepialidae, both represented by but few species and these rarely met with 

The Moths and Butterflies 


by collectors and nature students. But these moths are of particular impor- 
tance and interest to entomologists because they are undoubtedly the oldest 
or most generalized of living Lepidoptera; they represent most nearly, among 
present-day existing moths, the ancestral moth type. This is shown most 
conspicuously by the similarity in size, shape, and venation of the fore and 
hind wings, for the primitive winged insects had their two pairs of wings 
equal, while nowadays the various orders show a marked tendency to 
throw the flight function on one pair, either the fore wings, as among the 
flies (Diptera), wasps, bees, etc. (Hymenoptera), and Lepidoptera, or the 
hind wings, as with the locusts, crickets, etc. (Orthoptera), and beetles (Cple- 
optera), the other pair becoming much 
reduced in size, or even, as in the 
Diptera, wholly lost. Quite as impor- 
tant, if not more, although not so con- 
spicuous, as an evidence of the ancient 
character of the jugate moths, is the 
condition of the mouth-parts, certain 
species in the group having true biting 
mouth-parts, with well developed man- 
dibles, short lobe-like maxillae, and short, 
truly hp-like labium. All other moths 
and butterflies have the mouth-parts 
specialized for sucking, with the man- 
dibles rudimentary or wanting, the max- 
illje produced and apposed to form the 
long flexible sucking-tube, and the under 
lip (labium) reduced to a mere immovable 
functionless sclerite. The presence of 
the jugum for tying the fore and hind 
wings together, as in the caddis-flies. Fig. 524. — Diagram showing venation 
undoubtedly nearly alhed to the moth of wings in monarch butterfly, ^^mm 
■' ■' plexippus. c, costal vein; sc, sub- 

ancestors, instead of the specialized costal vein; r., radial vein; cm., cubi 

frenulum as in other moths, is also evi- 
dence of the ancestral type displayed by 
the Jugatae. 

The Micropterygidae, represented in 
this country by two genera, Eriocephala, 
with four species, and Epimartyria (Micropteryx) , with two species, are among 
the smallest moths we have, the largest not expanding more than one-third 
of an inch and the smallest only one-fifth of an inch, the body being about 
one-tenth of an inch long. They are indeed almost . invisible when flying, 
and are only very rarely taken by collectors. They fly in the sunshine, 

tal vein; a., anal veins. The base of 
the medial vein (lying between radius 
and cubitus) is obsolete, but its 
branches still persist, lying between 
branches of radius and cubitus. 
(Natural size.) 


The Moths and Butterflies 

frequenting flowers, and the different species are so much alike as to be 
nearly indistinguishable to the amateur. The eggs are laid on leaves, or in 
tiny pits in them, and the minute larvae, short and oblong, are either foot- 
less and mine the leaf substance, or have eight pairs of abdominal legs and 
feed exposed on leaves or in moss. The leaf-mining larvae burrow into the 
ground to pupate, while the exposed feeders make a slight cocoon of silk and 
debris above ground. The pupae are more like caddis-fly pupaj than the 
usual lepidopterous chrysalids (another indication of the primitive char- 
acter of the family), and those of certain species have large mandibles which 
they use to cut their way out of the cocoon. The adults can best be dis- 
tinguished by the venation of the wings (Fig. 525), and if ever found should 
be highly prized by the collector as specimens of the most primitive living 

The Hepialidae, the ghosts or swifts, although an offshoot from the 
Micropterygidae, or at least much more nearly related to them than to 
any other moths, are very different in appearance, being from an inch to 
2k inches long (some foreign species have a wing expanse of 6 inches) 
with large broad-ended wings and rather heavy body. They can be recog- 
nized by their venation (Fig. 526), which distinguishes them from all other 
moths of their size. The mouth-parts are rudimentary, but the parts per- 

sc rjr3r3r4 

~m — r. — '<^^r~p^mJ 

a a ^''' y 

Fig. 525. Fig. 526. 

Fig. 525. — Diagram of wing venation of Micropteryx sp. cs, costal vein; sc, subcostal 
vein; r, radial vein; m, medial vein; c, cubital vein; a, anal veins. (After Com- 
stock; enlarged.) 

Fig. 526. — Diagram of wings of Hepialus gracilis, showing jugum (7), and similarity of 
venation in fore and hind wings. (After Comstock.) 

sisting indicate plainly that they are reduced remnants of a very simple set 
of structures. The labium is free and truly lip-like and of the type of the 
under hp of biting insects. Two genera, Sthenopis, four species, and Hepi- 
alus, nine species, occur in this country. All of these moths are rather sombre 

The Moths and Butterflies 


in color, being grayish, yellowish brown, and reddish brown, with a few 

silvery-whitish irregular streaks on the upper 
wing surface. They fly swiftly and are said to 
prefer twilight. The males of some species give 
off a strong scent to attract the females. Others 
seem to show off their silvery spots by hovering 
for some time in the air at twilight, being con- 
spicuous, despite the semi-darkness and the 

^^^- ^^"^ll^-^^n '^'l"*«^s-moth, -g^ p-eneral coloration of the moth, by a pale 
I inea peUwnella; larva, larva ^ o j . r 

in case, and adult. (After silvery appearance. Females have been seen 
Howard and Marlatt; twice ^^ fly directly to the ghostly hovering males 

natural size.) .. ' , _,, . i-. , 

as if strongly attracted. Ihe grub-like larvae 
feed in the roots of various plants, as ferns and others, or in the trunk-wood 
of various shrubs and trees, and live for two or three years. Sthenopis 
argenteo-maadatus feeds first in the roots of alder, later going into the 
stems. It either pupates in its burrow or in a loose cocoon in the soil. 
The pupce are provided with certain short spiny teeth, and can wriggle so 
strongly that they are able to move about in the burrows or soil, and when 
ready to transform work their way to the surface of the ground. 

The Jugat£e are looked on by Comstock as equivalent in ranking to all 
the other moths and all the butterflies combined which are given the sub- 
ordinal name Frenatae. That is, this scant dozen of persisting represen- 
tatives of the ancient moth type, or rather 
of immediate offshoots from the ancestral 
type, are to be distinguished subordinally 
from all other living Lepidoptera, however 
more striking may appear the differences 
between some of these, as the obscure 
clothes-moths and the regal Cecropias, or the 
dull moth-millers and the brilliant day-fly- 
ing butterflies. The Frenate Lepidoptera 
include all those forms which have the vena- 
tion of the hind wings reduced (branches 
less in number than in the fore wings) 
and whose wings are tied together by a 
frenulum (Fig. 533) or by the expanded 
humeral angle of the hind wing overlapping 
the base of the ' fore wing, or by no more 
elaborate means than the simple overlapping 
of front margin of hind wing and hind 
margin of fore wing, but never by a jugum, 
the caddis-fly-like method common to the Micropterygids and Hepialids. 

Fig. 528. — Larva of the palmer- 
worm, Ypsolopliiis pomatellus, 
lying under its web spun on a 
leaf (After Lowe; natural length 
\ inch.) 


The Moths and Butterflies 

Among the Frenatae there is a host of small obscure moths commonly 
lumped by collectors and amateurs under the name Microlepidoptera, which 

are little known because httle 
studied, but which professional 
entomologists recognize as in- 
cluding all together eleven moth 
families grouped into three dis- 
tinct superfamiHes. Among these 
microlepidoptera are probably the 
most generahzed of the frenate 
Fig. 529. Fig. 530. moths. 

Fig. 529. — The palmer-worm moth, Ypsolophiis The three microlepidopteroUS 
P^-^^atellus. (After Fitch; twice natural s^perfamilies are the Tineina, 

Fig 530. — The strawberry root-borer> Anarsia including the clothes-moths, leaf- 
Uneatella. (After Saunders; moth and larva miners, and Others, the Tortri- 
both natural size and enlarged.) . 

cina, including most of the leaf- 
rollers, the notorious codlin-moth and others, and the Pyralidina, including 
certain leaf-rollers and folders, the close-wings, the curious plume-moths, the 
injurious meal-moths, and the bee-moth, principal pest of the bee-keeper. 

The Tmeidae, only family of the Tineina, are best known by their house- 
hold representatives, the clothes-moths. Of these there are several species, 
the moths themselves looking much alike, although distinguished by some 
differences in marking, but the larvae, the stage in which the injury to woolens, 
etc., is done having noticeable differences in habit. The moths lay their 
eggs on garments and stuffs, preferably woolen, hanging in dark closets or 
stored in trunks or dressers, and the small white larvae feed on the dry 
animal fibers of which the cloth is made. The larva of the most familiar 
species, the case-bearing clothes-moth. Tinea pellionella (Fig. 527), makes 
a small free tubular case out of bits of cloth fibers held together by silk'spun 
from its mouth; the larva of the tapestry -moth T. tapetzella, a rarer species, 
attacks thick woolen things, as blankets, carpets, and hangings, burrowing 
into the fabric and forming a long winding tunnel or gallery partially lined 
with silk; the larva of the webbing clothes-moth, Tinea biselliella, a species 
especially common in the Southern States, although not infrequent in the 
North, spins no case or gallery, but makes a cobweb covering over the 
substance it is feeding on. The larvae of all the species, when ready to 
pupate, make a cocoon out of bits of woolen tied together by silken threads 
in which to transform. The moths, on issuing, rest during the day on the 
garments or stuffs, but fly about at night, often coming to the lights in 
rooms. They are all small, pellionella and biselliella expanding about -j 
inch and tapetzella | inch; pellionella has grayish-yellow fore wings with- 
out spots, and tapetzella has the 'fore wings black at base and creamy- 

Z. p. mlz. I uALF* 

The Moths and Butterflies 


white with some grayish on the middle and apex. The eggs are laid 
by the moths directly on the woolen garments or other articles favored 
by the larval palate, and several generations may appear each year. The 
remedies for clothes-moths are the admission of light into closets and dressers, 
the fumigation of infested clothes or rugs in tight chests with bisulphide 
of carbon (the fumes will kill every larva and moth in the chest), and the 
keeping of carpets, rugs, hangings, and garments in cold storage during 
summer absences from home. Send the things to a 
cold-storage company with instructions to keep at 
a temperature below 40° F. The insects cannot 
develop in a temperature below this point. Cloth- 
covered furniture and cloth-lined carriages, if to be 
left long unused, may be sprayed once each in April, 
June, and August with benzine or naphtha. 

A sometimes serious pest of stored grains, espe- 
cially corn in cribs, is the Angoumois grain-moth, 
Gelechia cerealella. The larvae bore into the kernels, 
feeding on the inner starchy matter. I have seen ears 
of corn in Kansas cribs with every kernel attacked. 
The larvae feed for about three weeks, then pupate 
inside the kernel, the moth issuing in a few days. 
The kernels of infested ears show from one to 
three little holes from which moths have issued. 
The adult moth, expanding about half an inch, is 
light grayish brown, more or less spotted with black, 
looking much hke the case-bearing clothes-moth. 
The eggs are deposited on grain in the field or bin. 

Numerous Tineid species are known as leaf- 
miners because of the burrows of the larvae. Leaves 
of various trees and shrubs often show whitish blotches 
or lines, which when examined closely are seen to 
be due to the separation of the epidermis of the leaf 
from the inner soft tissue or to the complete dis- 
appearance of the inner tissue. This is the work of 
the tiny burrowing and feeding "leaf-miners," the 
larvae of certain Tineid species. Often the miner, 
a small white grub with the usual eight pairs of legs characteristic of Lepi- 
dopterous larvae, can be found in his mine, or, perhaps he will have ceased 
feeding and have transformed to a small light-brown pupa. The species of 
these leaf-miners are many, and numerous different types of mines may be 
found; the winding narrow lines called serpentine mines common on wild 
columbine, the spotted and folded tentiform mines on the wild cherry and the 

Fig. 531. — Pupal cocoons 
of the apple bucculatrix, 
Bucculatrix pomijoliella. 
(Twice natural size.) 


The Moths and Butterflies 

apple, the blotch-mines of the oaks and other forest trees. Even pine- 
needles are mined by certain species, the pine leaf-miner, Gelechia pini- 
jolieUa, being abundant in the leaves of pitch-pine. 

Interesting little Tineids are the apple and oak bucculatrix-moths, whose 
larvae feed on the leaves and w\\tn ready to pupate crawl to a stem or branch 

ff r2r.3 r4 

'r-\--\ M \ ^^ C2 

Fig. 532. . Fig. 533. 

Fig. 532. — The apple-leaf bucculatrix, Bucculatrix pomifoliella, pupal cocoons on twig, 
one pupal cocoon removed, and moth. (After Riley; cocoons natural size; 
size of moth indicated by line.) 

Fig. 533. — Venation of a Pyralid moth, P>'ra/:'5 /ar/;ja/f5. C5, costal vein; 5C, subcostal 
vein; r, radial vein; m, medial vein; c, cubital vein; a, anal veins. Note the hair- 
like projection, called frenulum, at the base of the anterior margin of the hind wing. 
This fits into a little "frenulum pocket" on the fore wing. (After Comstock; 

and there make long, slender, finely woven little white cocoons, conspicuously 
ribbed or fluted lengthwise, in which they pupate (Figs. 531 and 532). The 
pupae hibernate, the tiny moth issuing the following spring and laying its 
eggs on the leaves. The larva are miners at first, but after the first moulting 
feed on the outer surface of the leaves under thin flat silken webs. 

The PyraliJina include half a dozen famiUes, some of the moths 
hardly properly called microlepidoptera, for they reach a wing expanse of 
i^ inches. But most of the species are small and but few are at all 
familiar to collectors. The larvae of numerous species are injurious to 
fruits, stored grain, etc., and these species have a particular interest for 
economic entomologists. To collectors and nature students the most attrac- 
tive Pyralids will be the beautiful plume-moths, or feather-wings, small 
moths with the wings split or fissured longitudinally for one-half or more the 
length of the wing. The fore wings are usually thus divided into two parts 
and the hind wings into three (Fig. 534), but on some there are more divisions. 
All the feather-wings excepting one species belong to the family Pteropho- 

The Moths and Butterflies 


ridte, the exception being a small moth with both wings deeply cleft into 
six parts. It is called Orneodes hexadadyla and is considered to be the 
sole representative, so far as known, of a distinct family, the Orneodidai. 
Of the Pterophoridae several species are common in the North and East. 

Fig. 534. 

Fig- 535- 

Fig. 534. A California plume-moth. (Natural size.) 

Fig. 535. — The raspberry plume-moth, Oxyptilus tenuidactylus, moth and larva. (After 
Saunders; moth natural size; larva much enlarged.) 

Oxyptilus tenuidactylus (Fig. 535), with coppery brownish wings, with the 
plumes deeply fringed, has a pale yellowish-green larva that feeds on rasp- 
berries and blackberries; O. periscelidactylus has wings of a metallic yellow- 
ish brown, with several dull whitish streaks and spots; its greenish-yellow 
caterpillars with scattered small tufts of white hairs feed on grape-leaves 
and often are numerous enough to do much damage. Along the Pacific 
coast the plume-moths are not at all uncommon. 

Tig. 536. — ^The Mediterranean flour-moth, Ephestia kuehniella; larva, pupal cocoon, 
pupa, and moth. (One and one-half times natural size.) 

The Crambids, or close-wings, are numerous and perhaps more familiar 
than any other family of the Pyralidina. The larvae of most of the species 
feed on grass, and the adults fly up before one as one walks through meadow 
or pasture. They may easily be recognized by their characteristic habit of 
closely folding their wings about the body when at rest. The fore wings 
often present pretty designs in silver, gold, yellow, brown, black, and white, 


The Moths and Butterflies 

or they may be uniformly dull-colored; the hind wings are white or grayish. 
The palpi are long and project conspicuously, so that snout-moth is a name 
often given to the Crambids. 

Pretty little moths with shining black wings, two-spotted with white on 
the front ones, and one- or two-spotted on the hind wings, are the Desmias, 
of which the species maculalis, the grape-vine leaf-folder, is especially common, 
and often seriously injurious. The larvae fold or roll up grape-leaves and 
feed concealed inside the roll, skeletonizing the leaf by eating away all of its 
soft tissues. The larva when full-grown is a little less than an inch long, 
glossy yellowish green, and very active when disturbed. It pupates within 
the folded leaf. It is abundant in the South. 

Among the insects that attack stored grain, flour, meal, etc., are several 
Pyralids. The meal snout-moth, Pyralis jarinalis, is a common pest, 
the larvae making long tubes of silk in the meal, and taking readily to cereals 


Fig. 537. — A curious hammock and its maker, Coriscum ciiculipennellitm, a leaf-rolling 
moth, whose larva pupates in the odd little hammock shown in the figure. (After 
photographs by Slingerland; natural size of moth indicated by line; hammock 
natural size; a rose-leaf enlarged.) 

of all kinds and conditions, in the kernel or in the form of meal, bran, or 
straw. The moth expands one inch, the wings being light brown with red- 
dish reflections and a few wavy transverse lines. The Indian meal-moth, 
Plodia inter punctella, is another familiar pest in mills and stores, its small 
whitish larva, with brownish-yellow head, feeding on dry edibles of almost 
every kind, as meal, flour, bran, grain of all sorts, dried fruits, seeds, and nuts, 
condiments, roots, and herbs. It spins webs of silk with which it fastens 
together particles of the attacked food, making it unfit for our use. The moth 
expands f inch and has the fore wings cream-white at base and reddish 

The Moths and Butterflies 379 

brown with transverse blackish bands on disk and apex. Another and per- 
haps the most formidable of all mill pests is the notorious Mediterranean 
flour-moth, Ephestia kiiehniella (Fig. 536). This insect first became seri- 
ously harmful in Germany in 1877, soon invading Belgium and Holland 
and by 1886 having got a foothold in England. Three years later it 
appeared in Canada and since 1892 it has been a pest in the United States. 
The moth, which expands a little less than an inch, with pale leaden-gray 
fore wings, bearing zigzag black and transverse bands and semi-transparent 
dirty-whitish hind wings, lays its eggs where the hatching larvae can feed on 
flour, meal, bran, prepared cereal foods or grain. The caterpillars spin 
silken galleries as they move about, which make the flour lumpy and stringy 
and ruin it for use. In addition to this direct injury, the mill machinery 
often becomes clogged by the silk-filled flour and has to be frequently stopped 
and cleaned, involving in large mills much additional loss. When a mill 
becomes badly infested the whole building has to be thoroughly fumigated 
by carbon bisulphide, an expensive and rather dangerous process. Unin- 
fested mills should be tightly closed at night (if not running continuously) 
and every bushel of grain, every bag or sack brought into the mill, should 
be subjected to disinfection by heat or the fumes of bisulphide of carbon. 

An interesting as well as economically important little Pyralid is the 
bee-moth, Galleria tnellonella, whose larvae live in beehives, feeding on the 
wax combs. The moths find their way into the hives at night to lay their 
eggs. This has to be done very quickly, however, as bees are alert even at 
night to defend themselves against this insidious enemy. I have intro- 
duced bee-moths into glass-sided observation-hives both by day and night, 
and in each case the moths were almost immediately discovered, stung to 
death and torn to pieces in a wild frenzy of anger. Many must be killed 
where one succeeds in getting its eggs deposited inside the hive. The squirm- 
ing grub-like white larvae protect themselves by spinning silken webs and 
feed steadily on the wax, ruining brood- and food-cells and interfering sadly 
with the normal economy of the hive. When ready to pupate they spin 
very tough bee-proof silken cocoons within which they transform to other- 
wise defenceless quiescent pupai. Bee-moths often become so numerous 
in a hive as to break up the successful life of the community. I have taken 
thousands of pupae, lying side by side like mummies in sarcophagi in their 
impervious stiff silken cocoons, from a single hive from which the bees had 
all fled. 

Third of the superfamilies of microlepidoptera is the Tortricina, com- 
prising three families, two of which number many species. The Tortricid 
moths get their name from the habit, common to the larvae of many of them, 
of rolling up the edges or the whole of leaves in which to lie protected while 
feeding, and later while in quiescent pupal stage. Not all leaf-roUers are 


The Moths and Butterflies 

Tortricids, but the majority of rolled-up leaves so commonly seen on shrubs 
and trees are the homes of these larvae. A number of species belonging to 
the genus Cacoecia are among the commonest and most important of these 
because they prefer the leaves of apple, plum, and cherry trees, and currants, 
raspberries, gooseberries, strawberries, cranberries, roses, etc., rather than 

those of trees and shrubs 
-^'-^ — iif ^ whose healthfulness is not 

so important to us. The 
larvae of Cacoecia rosaceatia, 
the oblique-banded 1 e a f - 
roller, pale yellowish-green 
caterpillars | inch long, dis- 
figure and injure many kinds 
of fruit-trees, small fruits, 
and garden shrubs. The 
moth expands about one 
inch, and has reddish-brown 
body, light, cinnamon-brown 
fore wings crossed by wavy 
dark-brown lines and ochre- 
yellow hind wings. Choke- 
berries, and cultivated cher- 
ries as well, are often attacked 
by the cherry-tree leaf-folder, 
C. cerasivorana (Fig. 538), whose active yellow larvae "fasten together with 
silken threads all the leaves and twigs of a branch and feed upon them, 
an entire brood occupying a single nest. The larvae change to pupae within 
the nest; and the pupae when about to transform work their way out and 
hang suspended from the outer portion of the nest." The moths expand 
from -f to i^ inch, have bright ochre-yellow wings with brownish spots, and 
bands of pale leather-blue on the front ones. 

The oak leaf-roller, C. pervadana, similarly makes ugly nests in oak- 
trees in late summer, each nest consisting of a wad 
of tied-together leaves. Cranberry-plants are sometimes 
attacked by reddish, yellow-headed, warty-backed cater- 
pillars, which are the larvae of C. parallela (Fig. 540), 
a leaf-roller moth with reddish-orange fore wings crossed 
diagonally by numerous fine Hnes of a darker red-brown, 
and a pair of broad oblique red-brown bands. The hind 
wings are pale yellow. 

Notwithstanding the apparently sufficient protection afforded the leaf- 
rolling larvae by their tightly rolled cylindrical cases and webby nests, birds 

Fig. 540. 

Fig. 538. — The cherry-tree leaf-roller, Cacoecia cera- 
sivorana. (After Lugger; natural size.) 

Fig. 53Q. — Venation of a Tortricid, Cacoecia cera- 
sivorana. cs, costal vein; sc, subcostal vein; 
r, radial vein; m, medial vein; c, cubital vein; 
a, anal vein. (After Comstock; enlarged.) 

Fig. 540. — The cranberry leaf-roller, Cacoecia paral- 
lela. (After Lugger; natural size.) 

Fig. 541. — The 
tortrix, Dichelia 
siiljureana. (Af- 
ter Lugger; nat- 
ural size.) 

The Moths and Butterflies 


Fig. 542. — The rus- 
set-brown tortrix, 
Platynota flavedana. 
(After Lugger; 
natural size.) 

may often be seen cleverly engaged in extracting one by one the toothsome 

morsels from their homes. Hovering over a rolled leaf, the bill is carefully 

thrust into the roll for the unseen caterpillar and rarely vi^ithdrawn without 

it. Lugger says that the Baltimore oriole is particularly expert at this sort 

of hunting unseen prey. 

A certain Tortricid, accidentally imported many years ago from Europe, has 

become one of our serious grape pests. This is the grape-berry moth, Eu~ 

demis botrana, whose small slender whitish-green, black- 
headed larvx^ bore into green and ripening grapes and 

feed there on the pulp and seeds. When full-grown the 

larva becomes olive-green or dark brown and, forsaking 

the grape-berry, cuts out of a grape-leaf a little flap which 

it folds over and fastens with silk, thus forming a small 

oblong case within which it pupates. The moth expands 

finch, and has slaty-blue fore wings, marked with dark 

reddish-brown bands and spots, while the hind wings are uniform dull brown. 

Another well-known Tortricid pest is the bud-moth, Tmetocera ocellana 

(Fig. 543), whose larvae burrow into opening fruit- and leaf -buds on apple- 
trees and eat them. The moth expands f inch and is 
dark ashen-gray with a large irregular whitish band on 
the fore wing. 

By far the best known and most feared and hated 
Tortricid is the codlin-moth, Carpocapsa pomonella (Figs. 
545 and 546), the most important enemy of the apple- 
grower. Distributed all over the United States, wherever 
apples are grown, minute and obscure so as to be 
easily overlooked until fairly intrenched in the orchard, 

prolific and subject to no very disastrous parasitic attacks, this frail little 

species causes losses to fruit-growers of no less than $10,000,000 annually. 

The moth, which hides by day and is seldom seen, has the fore wings 

marked with alternate irregular transverse wavy streaks of ash-gray and 

brown, with a large tawny spot on the inner 

hind angle, the hind wings and abdomen 

light yellowish brown with a satiny luster. It 

lays its eggs (for the first generation, the species 

being two-brooded over most of the country) 

in the calyx of the newly forming apple, or 

sometimes, as recently observed in California, 

on the side of the tiny fruit. The larvae, hatch- 
ing in from three to five days, begin to feed on 

the green fruit, soon burrowing into its center. 

They become full-grown before the apples ripen, burrow out and crawl 

Fig. 543, — The eye- 
spotted bud-moth, 
Tmetocera ocellana. 
(After Lugger; 
natural size.) 

Fig. 544. — The cranberry 
worm-moth, Rhopotota vac- 
ciniana. (After Lugger; 
natural size indicated by 


The Moths and Butterflies 

away to some crevice in the bark or sheltered place on the ground, and 
there pupate. In two weeks the moths issue and deposit eggs on later 
apples for the second brood. The larvae of this brood are tucked away 
in the fall and winter apples when gathered, and are thus carried with 
them into cellars, warerooms, etc. They soon issue from the fruit, and 
finding concealed spots in the cracks of barrels or boxes or elsewhere 
near the stored apples, pupate, the pupae lasting over the winter and 
the moths issuing about apple-blossoming time the following spring. The 
pupae are protected by thin papery cocoons of silk spun by the larvae. The 
remedies are effective, but must be carefully and regularly used. Spraying 
the young fruit with an arsenical mixture, as Paris green or London purple, 
soon after the blossoms fall and again in about two weeks, will reduce 
immensely the possible loss. Banding the tree with strips of old carpet or 

Fig. 545. — The larva or worm of the codlin-moth, Carpocapsa pomonella. (After 
photograph by Shngerland; three times natural size.) 

sacking at the time the larvae are crawling out of the apples and hunting 
for concealed places in which to pupate, will enable the grower to trap and 
destroy thousands of them and thus greatly lessen the numbers in the second 
brood. All fallen fruit should be promptly gathered and destroyed in such 
a way as to kill the larvae inside. 

An interesting insect closely allied to the codlin-moth is the Mexican 
jumping bean-moth, Carpocapsa saltitans (Fig. 547), which lays its eggs 
on the green pods of a euphorbiaceous plant of the genus Croton. The 
hatchino- larvae bore into the growing beans in the pod, but do not attain 
their full growth until after the beans are ripe and hard. The ripe beans 
with the squirming larvae inside act as if bewitched, twitching and jerking, 
rolUng over and leaping shghtly clear of the table or desk on which they 

The Moths and Butterflies 


may rest. The larva; pupate within the beans, first gnawing a circuUu 
thin place through which the moth may push its way out. Another Tor- 
tricid moth, Grapholitha sebastiance, has similar habits. Most of the jump- 
ing beans come from the Mexican province of Chihuahua. 

Fig. 546. — Pupae, in cocoons, of codlin-moth, Carpocapsa pomonella. (After photograph 
by Slingerland; enlarged.) 

A few moth families, represented in this country by but few species, may 
now be referred to briefly, chiefly for the sake of mentioning certain par- 
ticular forms that are fairly common and wide-spread and hence likely to 
be taken by the collector. 

The flannel-moth family, Megalopygidae, includes but five North Ameri- 
can species, of which the crinkled flannel-moth, Lagoa crispata, pale straw- 
yellow, with long, curling, woolly, 
brownish and blackish hairs, with 
wing expanse of about i inch, is 
not uncommon in the north Atlantic 
states, while Megalopyge opercularis, 
of about the same size, with yel- 
lowish-white fore wings overspread 
except at the tips by woolly purpHsh- Fig. 547.— The Mexican jumping bean-moth, 
brown hairs, is not uncommon in C<^rpocapsa saltitans; pupa croton-bean 

from which moth has issued, and moth, 
the southern states. The flannel- (Natural size.) 

moth caterpillars have seven pairs 

of abdominal prop-legs instead of five, the number common to almost all 
other caterpillars, and the cocoons in which the pupae lie have a hinged 
door for the exit of the moth. The larva of M. opercularis looks like an 
animated bit of cotton-wool or lock of white hair. That of L. crispata 
feeds particularly on blackberry, raspberry, and apple; it is nearly oval 
in shape, covered with evenly shorn brownish hairs, which form a ridge 
along the middle of the back. When about \ inch long it ceases to feed 


The Moths and Butterflies 

and spins a tough oval cocoon fastened securely to the side of a twig. 
The moth issues in the summer of the following year. The cocoon of M. 
opercularis so closely resembles a terminal bud of the Southern live-oak on 
which the caterpillars mostly feed that it is almost impossible to detect it, 
especially as both twigs and cocoons are covered with small bits of lichen. 

Another small family, with thirty-three species, of interest because of 
the odd character of the larvae, is that of the slug-caterpillar moths, the 
Eucleidae (or Cochlidiida?). The moths themselves are small and stout, 
mostly rather strikingly colored, with brown, apple-green, and cinnamon 
prevailing. The larvte are slug-like, short, thick, nearly oblong and mostly 
spiny and gaudily colored. The spiny oak-slug, formidably armed with 
branching spines and common on oaks and willows in the east, is the larva 
of Euclea delphinii, a small, robust, deep-reddish-brown moth with bright 
green spots on the wings. The saddle-back caterpillar, Sibine (Empretia) 
stimidea', has a striking squarish green blotch on the back, with an oval pur- 
plish spot in the middle. It has branching spiny hairs, which affect some 
persons like nettles, producing severe inflammation. It feeds on many 
plants, on oak and other forest trees in the east, and often on corn in the 
west. The moth is lustrous seal- and chocolate-brown, with a few small 
white dots on the wings. Another slug-caterpillar is the pale apple-green 
larva, with dorsal brown blotch, of Prolimacodes (Eulimacodes) scapha, 
a stout wood-brown moth, expanding one inch, with a curved silvery line 
on each fore wing, behind which the wing surface is paler than in front. 
None of the species of this family has been found west of the Rocky Moun- 
tains except in Texas. Parasa chloris has the fore wings brown at base 
and outer margin and elsewhere apple-green; the hind wings are clayey 
yellow. Its larva is bright scarlet with four blue-black lines along the back 
and with stinging yellow tubercles. It feeds on cherry, apple, and rose. 
Euclea pcemdata has chocolate-brown fore wings with an irregular bright 
green elongate curving blotch, and the hind wings soft wood-brown. 

The most extraordinary species in this family of moths with strange 
larvae is the hag-moth, Phohetron pitheciutn, whose larva is one of the 
oddest known. It is nearly square, dark brown, and bears eight singular 
fleshy processes projecting from the sides. These processes, which are half 
as long as the larva itself, are covered with feathery brown hairs, among 
which are longer black, stinging hairs. Thus covered, and twisting curi- 
ously up and back, they resemble heavy locks of hair and give the name 
hag-moth to the species. The moth is rarely seen; it is dusky purple- 
brown with ocherous patches on the back and a light yellow tuft on each 
middle leg; the fore wings are variegated with pale yellowish brown, and 
crossed by a narrow wavy curved band of the same color; the hind wings 
are sable, bordered with yellowish in the female. 

The Moths and Butterflies 


Much larger moths are the Cossidaj, or carpenter-moths, with slender, 
smooth, spindle-shaped bodies and long, narrow-pointed, strong wings hke 
those of the hawk-moths (Sphingidtc). The larva) are wood-borers, bur- 
rowing about in the heart-wood of locust- and other shade-trees and also of 
apple-, pear-, and other fruit-trees. The moths are mostly gray, vaguely 
patterned with white and blackish, although a few are conspicuously black- 
and-white spotted. They have no proboscis and hence can take no food. 
The moths fly at night and lay their eggs on the bark of the trees, the hatch- 
ing, grub-like, naked larvae burrowing into the hard wood, where they Hve 
for from two to four years, when they make in their tunnel a thin cocoon 
of silk and chewed wood to 
pupate within. When ready 
to transform, the pupa 
wriggles along the tunnel 
to its opening, so that the 
issuing moth finds itself in 
free air. The locust-tree 
carpenter-moth, Prionoxys- 
tus robinm (Fig. 549), or 
goat-moth, so called from 
its curious offensive odor, 
expanding i^ inches (males) 
to 2\ inches (females), has 
gray wings with irregular 
black lines and spots in 
the female, and darker 
fore wings and yellowish 
hind wings in the male. 
Its larvce feed on locust- 
trees and are often abun- 
dant enough to do much injury. The wood leopard-moth, Zeiizera pyrina, 
is strikingly spotted with black on a white ground color, and is common in 
certain eastern cities, its larvae infesting maples and other shade-trees. On 
' the Pacific coast the poplar carpenter-moth, Cossus popidi, with whitish 
fore wings shaded all over with blackish and irregular black Hnes, and hind 
wings yellowish gray, growing darker at the outer margin, is common, its 
larvae infesting poplars and cottonwoods. There are only twenty species 
in North America belonging to this family. 

FamiUar curiosities of entomology are the moving bags of silk and bits 
of twigs and needles occasionally found in cedars, firs, and arbor vitae. The 
"worms" which make these bags and carry them around, with all the body 
inside except the projecting head and thoracic legs, are the larvae of the 

Fig. 548. — Venation of a Cossid, Prionoxystus robinia. 
cs, costal vein; sc, subcostal vein; r, radial vein; 
m, medial vein; c, cubital vein; a, anal veins. (After 
Comstock; enlarged.) 


The Moths and Butterflies 

bag- worm moth, Thyridopteryx ephemera; for mis (Fig. 550), the females of 
which are wingless, the males with blackish body and clear brown-veined 
wings which expand an inch. This moth is the most common and wide- 
spread of the thirteen moth species which constitute the family Psychids, as 
represented in this country. In the Southern States a common species is 
Abbott's bag-worm, Oiketicus abbotti, whose larvae make bags with the bits 
of twigs fastened regularly transversely, the male moth expanding ij inches 
and being sable-brown with a clear bar in the middle of each fore wing. 
Smaller bag-worm moths are the three species of the genus Psyche, the males 
expanding from ^ inch to | inch, P. conjederata, the best known, being all 

Fig. 549. — The locust-tree carpenter-moth, Prionoxystus robittics, male and female 
moths, young larva and empty pupal case. (After Lugger; moths and pupal case 
natural size; young larva enlarged.) 

blackish with opaque wings, P. gloveri, a Southern species, dark brown through- 
out, and P. carbonaria, a Texas form, brownish black with subtranslucent 
wings. The females of all the Psychids are wingless. The larvae, after 
moving about over the tree and feeding until full-grown, pupate within their 
bags, and the issuing wingless grub-like females simply remain in the sac 
until found by a flying male, after which they lay their eggs in the bag and 
die. The male Psychids can be readily distinguished from other moths by 
the growing together of the anal veins of the fore wings until they appear 
to be a single branching vein (Fig. 552). 

The smoky-moths, Pyromorphidae, of which but fifteen species occur , 
in the United States, are small, expanding from -| inch to i inch (a single 
Western species expands ij inches), and with blackish ground-color on body 

The Moths and Butterflies 


and wings, relieved by brilliant patches of red, yellow, and orange. They 
are favorites with collectors and, though few in number, are not at all uncom- 
mon. The larvai feed on the leaves of various plants, but grape and Vir- 
ginia creeper seem to be specially liked. Vineyards indeed often suffer 
from the presence in considerable numbers of smoky-moth caterpillars. 
These caterpillars often show a striking gregarious instinct, massing side 
by side in lines while feeding. The small black and yellow larvae of Har- 
risina americana, a common Eastern species, may often be found arranged 

Pig. 550. — The bag-worm moth, Thyridopteryx ephemercejormis; eggs, larva, pupa 
bag containing larva, bag containing pupa, male moth. (After Felt; about natural 
size except the eggs.) 

side by side in single line clear across a grape-leaf. Feeding, when young, 
only on the soft tissues of the leaves, they skeletonize them; when older, 
however, they eat everything but the larger veins. When full-grown they 
disperse, each finding a sheltered spot, where it makes a tough, oblong- 
oval cocoon of parchment-like silk, in which it pupates. The moth of this 
species expands one inch, is bluish or greenish black, with orange protho- 
racic collar broad above and narrow below, and narrow subtranslucent 
wings. It flies slowly and unevenly during the warmest, brightest hours 
of the day, frequenting flowers. H. coracina, found in Texas and Arizona, 
expands | inch and is all dull black with a bluish tinge on the abdomen; H. 
metallica^ the largest Pyromorphid, found in Texas and Arizona, expands 
1 1 inches and is lustrous bluish green with orange prothorax. Acoloithus 

The Moths and Butterflies 

jaJsariiis, one of the smallest members of the family, expanding f inch, com- 
mon in the East, is black with very narrow reddish collar. Pyromorpha 
dimidiaUi, expanding i inch, common in the Atlantic states, is black with 
translucent wings. The only other genus in the family so far unmentioned 

is Triprocris with eight species, all 
confined to the western states and all 
but two of them marked on body or 
wings with orange or yellowish. 

Of unusual and often very deceptive 
appearance are the clear-wing moths, 
or Sesiidae. With their often brightly 
colored black and yellow or red- 
banded tapering or plump bodies and 
partly or wholly clear wings, they 
resemble strongly, at first glance, wasps 
or bees, and are undoubtedly often 
taken to be such and thus left unmo- 
lested by both collectors and birds, two 
of their destructive enemies. For birds 
like almost all moths for food, and 
collectors especially prize the Sesians 
for the sake of their attractiveness 
and the sporting character of their pur- 
suit and capture, for they are among 
the swiftest of the moths. They fly 
in bright sunlight, visiting flowers, 
and thus by their habits further in- 
crease their likeness to wasps and bees. 
There are one hundred species in the 
family in this country, and almost all 
have one or both wings partly or mostly 
clear, i.e., free from scales. A few 
moths of other families, as the clear- 
winged sphinges and others, have simi- 
larly partly clear wings, but the very 
narrow fore wings and widely expanded 
bases of the hind wings will distinguish the Sesians from the few other 
scattered clear-winged moths. The larvae are borers, mining in roots of 
fruit-trees, the canes and roots of small fruits, or in the stems of herbaceous 
plants. They are grub-like and yellowish white, with darker head and legs. 
When abundant they become very injurious, the notorious peach-tree borers 
being probably the most serious insect enemy of the peach-tree. 

Fig. 551. — Bag-worm ; the larva of a moth 
that builds a protecting case out of 
silk and bits of sticks, in which its 
whole body except homy head, thorax, 
and legs is concealed. (Natural size-) 

The Moths and Butterflies 




For one hundred and fifty years the peach, an imported plant, has suffered 
in this country from the ravages of this native pest. One Sesian species, 
Sanninoidea exitiosa (Fig. 554) is 
the peach-tree borer of the eastern 
states, and another, closely related, 
S. pacifica, works equal injury in the 
Pacific states. In both species the 
eggs (Fig. 555) are deposited on 
the trunk of the tree near its base, 
in July and August in the East, in 
April and May in California, and 
the young larva; (Fig. 556), hatching 
after a week or ten days, immedi- 
ately bore in through the outer 
bark and begin feeding on the 
live inner bark. When winter 
comes they cease feeding — in the 
East a. Ieas,-and hibernate quies- ^'yil^ZTlt"Ti;T4!:^:'tf:- 
cent, being now about half-grown. merajormis. cs, costal vein; sc, subcostal 
In the spring they become active ^^in; r, radial vein; m, medial vein; c, 

^ ° -^ cubital vein; a, anal veins, 

again, feed and grow rapidly, and 

by summer are ready to pupate. Pacifica begins pupating in California 

in February. For this they leave 
their burrows, come out to the 
surface of the bark, spin about 
themselves a thin silken cocoon 
and change (Fig. 557). The 
pupal stage lasts 
weeks, when the 
The clear-winged 
expanding i inch, are deep 
steely-blue, with small golden- 
yellow markings on head and 
thorax and abdomen; the larger, 
heavier-bodied female, expanding 
a a a "* i|inches,has a broad orange band 

Fig. 553. —Venation of a Pyromorphid, Pyro- across the abdomen in the fourth 
morpha dimidiata. cs, costal vein; 5C, sub- q^. f^^^]^ segments, and has the 
costal vein; r radial vein; m, medial vein; . , . , , , 1 ■ 1 

c, cubital vein; a, anal veins. (After Com- front wmgs covered With blackish 
stock; enlarged.) scales (Fig. 554). The remedy 

for this pest is the application, by painting on, of gas tar to the basal 
part of the tree-trunk just before the flying and egg-laying time of the 

about three 
moths issue, 
male moths, 


The Moths and Butterflies 

moths; this prevents the females from ovipositing on the treated trees. Or 
the base of the trunk may have a newspaper tied about it. 

Fig. 554. — Moths of the peach-tree borer, Sanninoidea exitiosa, the upper one and the 
one at the right being females. (Photograph from hfe by Shngerland; natural size.) 

The currant-borer, Sesia tipuliformis, expanding three-fourths of an 
inch, has a robust body with a fan-hke tuft of scales at the posterior tip, 

Fig. 555. — Eggs of peach-tree borer, Sanninoidea exitiosa. (After Slingerland; natural 
size at n; one egg enlarged at I; micropyle end of egg greatly enlarged at m.) 

dark abdomen ringed with yellow, and yellow lines on the thorax; the eggs 
are laid on currant-canes, and the hatching larvae burrow into the center 
and then tunnel longitudinally in the pith. They hibernate in the cane 
as larvae, not pupating until ihe following summer, when the moths escape 

The Moths and Butterflies 


through holes in the cane thoughtfully made by the strong-jawed larvae 
before pupation. The grape-vine-root borer, Memythrus poUstijormis, looks 
much like a large Polistes wasp, having a dark body with two bright* yellow 

Fig. 556. — Larva of peach-tree borer, 5aKwmo/(/ea exitiosa. (After Slingerland; natural 

size and much enlarged.) 

narrow bands about the abdomen; the fore wings are brownish black, the 
hind wings clear; the larvae bore in the roots of wild and cultivated grapes 
and pupate underground. The raspberry-root borer, Bemhecia tnarginata, 
is also very waspish in appearance, with its black body repeatedly banded 


















Fig. 557. — Cocoons and empty pupal skins of the peach-tree borer, Sannmoidea exitiosa. 
(After Slingerland; natural size.) 

with yellow and transparent fore and hind wings. The eggs are laid on 
raspberry canes, and the larvae, first boring into the cane, finally work down 
into the roots. Squashes are often badly injured by having their stems 
tunneled by the larvae of the squash-vine borer, Melittia ceto, a Sesian with 
olive-brown fore wings, clear hind wings, and black or bronze abdomen, 


The Moths and Butterflies 

marked with red or orange, and with the hind legs fringed with long hairs, 
orange on the outer surface and black on the inner. When full grown the 
larvae leave the stems and go into the soil to cocoon and pupate. The genus 

Fig. 558. 

Fig. 559. 

Fig. 558. — The ash-tree borer, Trochiliiim fraxini. (After Lugger; natural size.) 
Fig. 559. — Sesia pidipes, male. (After Lugger; natural size.) 

Sesia (Fig. 563) contains over half (fifty-seven) of the species in this family; 

they are found in all parts of the country. 

The family Notodontidse, comprising the puss-moths, handmaid-moths, 

and prominents, is represented in 
this country by about ninety-five 
species, all of medium siz.e, i.e., with 
a wing expanse of from ij to 2 
inches, and but few of such marked 
patterns as to be particularly con- 
spicuous or attractive to collectors. 
The name " prominents," sometimes 
applied collectively to the moths of 
this family, is based on the occur- 
rence in some of them of an angu- 
lated or tooth-like projection near 
the middle of the hinder margin of 
the fore wings. Probably the most 
famiUar species in this family are 
the Datanas, or handmaid-moths; 
certainly their larvae are more often 
seen and are better known, under 
the names of yellow-necked apple- 
tree caterpillars and walnut cater- 
pillars, than the larvae of any other 

Notodontids. Sometimes there may be seen on the trunk of an apple- or 

other shade-tree an animated bunch or mass of hundreds of caterpillars, 

Fig. 560. — Venation of a Notodontid, Noto- 
donta stragula. cs, costal vein; sc, sub- 
costal vein; r, radial vein; m, medial 
vein; c, cubital vein; a, anal veins. 
(After Comstock; enlarged.) 

The Moths and Butterflies 


reddish black with conspicuous yellow longitudinal stripes, each caterpillar 

curiously jerking its body or resting quietly with both head and body tip 

held up nearly at right angles to the middle part with its four pairs of clinging 

prop-legs. These are Datana larvie, which 

have come down from their feeding on the 

leaves of the tree to moult. The jerking^,,. 

frightens away in some measure the numerous 

parasitic Tachina flies which are always 

ready to attend on a gathering of this sort 

and lay a few eggs where they will do the 

Tachina species the most good, that is, on 

the body of these plump caterpillars, so 

that the hatching Tachina grub can burrow into this well-nourished Ijody 

and feed on its living tissues. When feeding in the tree-tops, too, the Datana 

Fig. 561. — The red-humped cater- 
pillar-moth, CEdemasia eximia^ 
(After Packard; natural size.) 

Fig. 562. — Larva of red-humped caierpillar-moth, CEdemasia eximia. (After Packard; 

natural size.) 

caterpillars keep closely together, forming rows or files of voracious feeders 
arranged neatly across each attacked leaf. The common species infesting 
the apple is Datana ministra, and the larvce have a distinguishing dull orange 

spot on the back of the first body-ring 
behind the head. The eggs, which are 
white and spherical, are laid, from 70 to 
100 by each female, on the leaves, all 
cemented well together in neat patches. 
When the larvae are full grown they 
descend from the tree, burrow into the 
soil for two or three inches, and change 
to naked brown chrysalids, which last 
over winter, the moths emerging in the following summer. The moth, 
expanding ih inches, is reddish or yellowish brown, with the fore wings 
crossed by from three to five darker brown lines, the outer margin and one 

Fig. 563. — Heterocampa guttivitta. 
(After Packard; natural size.) 

394 The Moths and Butterflies 

or two spots near the middle also being darker; the hind wings are pale 
yellow and not patterned. The species common on walnuts and hickories is 
Datana angusit, with fore wings varying from chocolate to deep smoky 
brown, with transverse lines like those of ministra; the hind wings are 

Fig. 564. — Larva of Heterocampa guttivitta. (After Packard; natural size.) 

paler brown. The caterpillars are black, with dirty-white hairs and with 
three equidistant, very narrow, pale-yellow or whitish stripes on each side 
and three yellow stripes on the under side; when full grown it is a little more 
than 2 inches long. 

Another conspicuous Notodontid larva occurring on apple-trees is a 
greenish-yellow black-striped caterpillar with a coral-red head and promi- 
nent hump on the back of the fourth body-ring. This is the larva (Fig. 562) 
of the red-humped caterpillar-moth, (Edemasia concinna (Fig. 561), a 
darkish-brown moth expanding about i\ inches, the fore wings having a 
darker brown spot near the middle, a spot near each angle, and several 
longitudinal streaks along the hinder margin. 

The puss-moths, Cerura, are readily distinguishable by their characteristic 
black and white wings, white being the ground color, with two broad, not 
sharply defined blackish bars across the fore wing, one across the disk, the 
other, often incomplete posteriorly, across the apex. Along the outer margin 
of each wing there is a row of distinct small black points. The larvae (Fig. 
793) of Cerura are extraordinary creatures: short, thick, naked body, tapering 
behind to a kind of forked tail which is held up at an angle with the rest 
of the body. This tail, which is an organ of defence, consists of two tubes, 
within each of which is concealed a long orange-colored extensile thread 
which can be thrust out and drawn in at will. When disturbed, the puss- 
moth caterpillar thrusts out these vivid tails, waving them threateningly, 
at the same time giving off a strong odor. It also telescopes its head and 
front two thoracic segments into the large, humped, third segment, which is 
so shaped and marked as to suggest some formidable large-eyed creature 
quite unlike a soft-bodied toothsome caterpillar. With little doubt this 
elaborate terrifying but actually harmless equipment avails to frighten off 
many of Cerura's enemies. The larva of a common puss-moth species 
feeds on wild cherry. When ready to pupate the caterpillars gnaw out a 
shallow cavity or depression in the wood which they lie in and over which 
they spin an oval silken net mixed with particles of wood, which makes it 
almost indistinguishable from the rest of the wood surface. These moths 

The Moths and Butterflies 


seem to carry very far expedients of Nature for protection by deceit. Other 
common members of the family are the several species of Schizura, moths 
strongly resembling owlet-moths (Noctuidas) with their brown and gray 
and gray and blackish finely variegated fore wings and unmarked silky white 
wings. Their brown or greenish larvae, which feed on fruit-trees, forest 
trees, small fruits, and other shrubby plants, are distinguished by having 
a prominent horn or spined tubercle on the fourth body-ring behind the 
head. They are said to eat out a notch about the size of the body, in the edge 
of a leaf, fitting themselves along this notch, so that the prominent tubercle 
and other irregularities of the body seem to simulate the rounded edge of 
the leaf; they are thus well concealed. The moths, too, are much given 
to dissimulation. Each moth rests on the trunk or branches of the tree, 

Fig. 565. — Canker-worms, larvae of a geometrid moth. (After Slingerland; natural size.) 

head downward, with wings closely folded around the body and legs all 
drawn together, the dull-gray tone of the wings with their bits of lichen- 
green and whitish color giving the whole a marvelous resemblance to a bit 
of rough weathered bark. 

Familiar to all observers, although certainly not very often seen and 
rarely found in large numbers, are the inchworms, spanworms, or loopers 


The Moths and Butterflies 

Fig. 566. — Lime-tree inch-worm, larva 
of the geometrid moth, Hibernia 
tiliaria. (After Pettit; twice natural 


as they are variously called, which are the larvae (caterpillars) (Fig. 565) 
of the moths of the superfamily Geometrina (earth-measurers). These 
three common names as well as the scientific one refer to the peculiar mode 
of locomotion affected by all the Geometrina. Each loop or step is made by 
the bringing forward of the caudal extremity of the body quite to the thoracic 
feet, the portion of flexible body between 
bending up and out of the way each time 
during the process. The reason for it 
all will be understood when the inch- 
worm is examined. It differs from other 
lepidopterous larvae in lacking the front 
three of the four pairs of prop-legs 
normally belonging to the middle part 
of the body, which is thus rendered 

helpless in walking, and the curious looping gait is the outcome of the pos- 
session by a long slender flexible body of only anterior and posterior locomotory 
organs (Fig. 566). Why inchworms are not more often seen, although there 

are hosts of different kinds of them and they 
are well distributed and common all over the 
country, is due to their habit of "going 
stiff" when disturbed, clinging by the hinder 
two pairs of legs to the twig or leaf and 
holding the rest of the body motionless and 
rigid at an angle with the support. As the 
body is always protectively colored and 
marked, so as to harmonize thoroughly with 
the habitual surroundings many an inch- 
worm may be seen but not distinguished 
from the leaf or branch on which it rests. 
Indeed, many of the inchworms are amaz- 
ingly like a short or broken twig, with buds 
or leaf scars and lined or scaly bark, a very 
effective case of protective resemblance. 

The geometer-moths, of which we have 
800 species in this country, while of course 
presenting a great variety of coloration and 
pattern yet possess a likeness of general 
appearance due mostly to the slenderness of 
body compared with the broadness of wings, the impression of fragility or thin- 
ness of wings due to the unusually fineness of the covering scales, and the deli- 
cate and quiet coloration and patterning, which indicate their identity pretty 
effectively. Some are small, i.e., less than i inch expanse, and a few large, 

Fig. 567. — -Venation of a geometrid, 
Dyspepteris abortivaria. cs, cos- 
tal vein; sc, subcostal vein; r, 
radial vein; m, medial vein; 
c, cubital vein; a, anal veins. 
(After Comstock; enlarged.) 

The Moths and Butterflies 397 

i.e., over 2 inches expanse, but most are of medium size, with white, deli- 
cate green, soft yellowish, brownish, grayish, and blackish as predominating 
color tones, and delicate wavy or zigzagging transverse lines, or point-like 
spots as characteristic pattern markings. The superfamily is divided into 
five famiUes based on venational characters rather confusing and appar- 
ently not surely indicative of natural relationships. We may content our- 


Fig. 568. — Male and IriiKilc lime-tree canker-moths, Hibernia tiliaria. (After Jordan 
and Kellogg; twice natural size.) 

selves with brief reference to some of the more interesting, beautiful, or eco- 
nomically important species. 

The best-known Geometrids of economic importance are the canker- 
worms (Fig. 565), two species in particular, known as the spring canker- 
worm (Paleacrita vernata) and the fall canker-worm {Anisopteryx pometaria), 
being responsible for much damage to orchards, especially apple-orchards. 
The females of the canker-worm moths are wingless and so have to climb 
the trees to lay their eggs on the branches and twigs. 
This fact naturally suggests the most effective remedy 
for them, namely, banding the trees with tar (mixed 
with oil to prevent its drying) so as to make effective 
barriers against them as they crawl upward. Printers' „ ^ r^ . .. • 

° y r YlG. 50Q. — Dyspepterts 

ink, refuse sorghum, or any slow-drying varnish is abortivaria. (After 
equally effective. From the eggs laid in the spring by Lugger; natural 
Paleacrita and in the fall by Anisopteryx hatch active 
little "loopers" which feed voraciously in the foliage. The eggs of the fall 
canker-worm do not hatch until the following spring, just when the young 
apple-leaves begin to unfold. The full-grown canker-worms are about i 
inch long, greenish brown and striped longitudinally with pale yellow. 
Some of these stripes are broad on the fall canker-worm; all are narrow 
on the other species. When full grown the larvae crawl down the tree to the 
ground, burrow into it and pupate in a thin silver cocoon. The males of both 
species are winged delicate moths; Paleacrita has pale ash-colored or brownish- 
gray, silky, almost transparent fore wings with four or five broken transverse 


The Moths and Butterflies 

dark lines; Anisopteryx has glossy brownish fore wings crossed by two 
irregular whitish bands. 

Among the Geometrids are numerous species whose wings are green, 
the shades varying, but usually with a strong admixture of whitish and also 

Fig. 570. . Fig. 571. 

Fig. 570. — The pepper-and-salt currant-moth, Eubyia cognataria. (After Packard; 

natural size.) 
Yic. ;;7i. — Phigalia strigataria, the iemale wingless. (After Lugger; natural size.) 

usually barred more or less distinctly with narrow or broader whitish lines. 
Geometra iridaria is such a species common in the East in which the green 
is very light in tone; Dyspepteris abortivaria (Fig. 569) is bluish green and 

Fig. 572. Fig. 573. Fig. 574. 

Fig. 572. — The large blue-striped looper, Biston ypsilon. (After Forbes; natural size.) 

Fig. 573. — The common Cymatophora, Cymatophora pampinaria. (After Lugger; 

natural size.) 

Fig. S74. — The plum-geometer, Eumacaria hrunneraria. (After Lugger; natural size.) 

has a grape-feeding larva. The raspberry geometer, Synchlora glaucaria, 
has delicate pale-green wings with two transverse whitish lines; its larvse 
feed in the fruit and leaves of raspberries and blackberries and cover over 

the body with bits of vegetable matter like minute 
pieces of flowers, etc., until it seems to be only a 
tiny heap of debris. The snow-white Eugonia, 
Ennonos suhsignarius, is pure white, expanding 
an inch and a half; its larvae feed often de- 
structively on the foliage of elms, lindens, and 
apple-trees. Angerona crocotaria (Fig. 576) is 
a beautiful sulphur - yellow Geometrid, ex- 
panding i\ inches, with a number of irregular pinkish-brown blotches 
on the wings; its yellowish-green larvae feed on currants, gooseberries. 

Fig. 575. — The currant fruit- 
worm moth, Enpithecia in- 
trrruptofasciata. (After 
Lugger; natural size.) 

The Moths and Butterflies 


and strawberries, both wild and cultivated. Calocalpe undulata (Fig. 578), 
the scallop-shell moth, has pale yellowish-brown wings crossed by many 
fine zigzag darker lines close together; its larva? feed on wild cherry and 
live gregariously inside of a nest formed of leaves tied together by silken 
threads. A very common little moth in meadows and gardens in summer 
and fall is the chickweed-geometer, HcBmatopis grataria, with reddish- 

FiG. 576. Fig. 577. Fig. 578. 

Fig. 576. — The currant-angerona, Angerona crocataria. (After Lugger; natural size.) 
Fig. 577. — The currant-endropia, Endropia armaiaria. (After Lugger; natural size.) 
Fig. 578. — The scallop-shell geometer, Ca/oca//)e M«(fj</ato. (After Lugger; natural size.) 

yellow wings and two transverse bands and the outer margins pinkish 
The chain-dotted geometer, Caterva catenaria, expanding ij inches, with 
white wings dotted with fine black points arranged in two lines and with 
a few extra ones, appears sometimes, according to Lugger, in such very 
great numbers as to look like a snow-storm; its larvae are pale straw-yellow 
with two fine lines on the back and two on each side interrupted by two 

Fig. 579. — The diverse-lined geometer, Petrophora diver silineata. (After Lugger; 

natural size.) 

large black dots, a pair on each segment; it feeds on hazel, blackberry, 
raspberry, and other plants. 

A great host of somber-colored moths, blackish, grayish, or brownish, 
with no conspicuous markings and only rarely any bright colors, compose 
for the most part the family Noctuidse, the largest of all the families of moths. 
Twenty-one hundred North American species — three times as many as 
there are North American species of birds — belong to the single family 
Noctuidae, and for the most part these two thousand mixed species must be 
as one to the general collector and amateur. Few professional entomologists, 
indeed, lay claim to a systematic knowledge of the group, or even care to 
give to it the time necessary to acquire such a knowledge. Some of the 


The Moths and Butterflies 

Noctuids have come into prominence because of the destructive vegetable- 
feeding habits of their larva' ; such are the cutworm-moths, the army- 
worm moths, the cotton-worm moths, and others, and these species are 
so often described and pictured that they are fairly well known. Other 
small groups, of which the interesting Catocalas, the red and yellow under- 
wings (Fig. 580), are the most conspicuous, have attracted the attention 
of collectors because of particular habits or patterns, and these are fairly 

Fig. 580. — A group of red and yellow underwings; upper moth, Catocala palceogiima; 
lower left-hand corner, Catocala iiltronia; lower right-hand corner, Catocala grymea. 
(After Lugger; natural size.) 

well known. Few moth-collectors but have "sugared" for Catocalas, 
those large night-flyers, somber of fore wing but brilliant of hind wing, 
that can be so readily attracted and taken by a bait of molasses and stale 
beer smeared in patches on the trunks of trees in summer-time. The fore 
wings harmonize in color, shades, and pattern so thoroughly with the bark 
that when the Catocala rests, as it does during the daytime, on tree-trunks 
with its brilliant hind wings, strikingly banded with red, yellow, white, or 
black, covered by the fore wings, it is simply indistinguishable. The 
Catocala larvae are curious creatures, with body thick in the middle and 

The Moths and Butterflies 


tapering towards both ends. The lar\ie of Catocala nltronia (Fig. 581) 
feed on plum-tree leaves; tlicy arc about ij inches long, grayish brown, 
with two or four small reddish tubercles on each body-segment, a small 
fleshy horn on the back of the ninth segment and on the back of the twelfth 
segment a low fleshy ridge tinted behind with reddish brown. It descends 
to the ground when ready to pupate, making a flimsy cocoon of silk under 
a dead leaf or chip. The pupa inside the cocoon is covered with a bluish 
flour-like dust or "bloom." The moth has the forewings rich amber with 
a broad indefinite ashy band along the middle and several brown and 

Fig. 581. — The plum-tree Catocala, Catocala iiUronia, moth and larva. 
(After Lugger; natural size.) 

white transverse lines; the hind wings are deep red with a wide black 
band along the outer margin and a narrower one across the middle. The 
eggs are laid in cracks of the bark in slimmer. Catocala grynea (Fig. 580), 
with grayish brown forewings marked with zigzag lines of rich brown and 
gray short dark-brown streaks on the front margin and with hind wings 
reddish yellow crossed by two wavy black bands, is called the apple-tree 
Catocala, because the ashen-brown caterpillar feeds on apple-leaves. The 
two front pairs of abdominal prop-legs of all the Catocala caterpillars are 
much smaller than the hinder two pairs, hence the caterpillar has a sort of 
looping gait like that of the Geometrid larvae, the inchworms. Catocala 
relicta has the fore wings grayish white with several indefinite transverse 
black bands, and the hind wings black with one curving white band. 
Catocala epione has blackish-brown fore wings with wavy narrow black and 
lighter brown transverse lines with black hind wings narrowly rr^argined 
with white. 

The largest and most interesting Noctuid, and indeed one of the largest 
of all the moths, is the curious rare species Erebus odora, called the black 
witch; it expands 6 inches and has both wings blackish brown with many 


The Moths and Butterflies 

indefinite wavy lines of black and of lighter brown; in the hinder angle 
of the hind wings are two incomplete eye-spots bounded in front by a curv- 
ing velvety black line, and on each fore wing is a single irregular eye-spot 
near the front margin. 

"Cutworm" is the name applied to the smooth, "greasy," plump cater- 
pillars of numerous species (representing several genera) of Noctuids. The 
greasy cutworm, dull blackish brown with pale longitudinal Unes attacks 
all sorts of garden products and other low-growing plants; it is the larva 

Fig. 582. — Green-fruit worms, Xylina grotci, at left, and Xylina antennata at right. 
(Photograph by Shngerland; natural size.) 

of A gratis ypsilon, with brownish-gray fore wings bearing an ypsilon- 
shaped mark, the hind wings being silky white. The climbing cutworm, 
Carneades scandens, an active climber and great enemy of nurseries and 
orchards, is light yellowish gray with a dark line along the back and fainter 
ones along the sides; the moth has light bluish-gray fore wings with darker 
markings and pearly-white hind wings. Almost all the cutworms hide 
in cracks in the ground by day, feeding during the night; they will often 
cut off young plants just at the ground, or will ascend tall trees and feed 
on the buds and young leaves. When ready to pupate they burrow into 
the soil and the moths issue in midsummer. 

The. members of the large genus Plusia (PI. VIII, Fig. 7), including some 
of the commonest Noctuids, are recognizable by a small silvery comma-shaped 
spot on the disk of each fore wing. Another large genus is that of Cucul- 
lia, the hooded owlets, in which the thorax bears a prominent tuft of scales 
and the fore wings are marked with irregular blackish dashes. The 

The Moths and Butterflies 


Fig. 583. — Army-worms, larvae of Leucania unipuncta, on corn. (Photograph by 
Slingerland; natural size.) 


The Moths and Butterflies 

dagger-moth Acronycta (Figs. 586 and 587), so called from the rather uncer- 
tain small black dagger-like markings of the fore wings, have the larva in 
some species covered with long colored stiff hairs; the familiar caterpillar 

of A. americana is densely clothed with 
yellow hairs, besides bearing a pair of 
long black pencils on the first abdominal 
segment, another pair on the third, and 
a single pencil on the eighth. It feeds on 
the leaves of elm, maple, and other trees, 
and when at rest curls sidewise on a leaf. 
The army- worm (Fig. 583), a black, 
yellow, and green striped caterpillar 
that occurs over nearly all the country 
and often appears in enormous numbers, 
causing great losses to grain-fields, is 
the larva of a dull-brown moth, Leu- 
cania unipuncta, marked in the center 

costal vein-^W °^ ^^^'^ ^°^^ ^^"S ^^^^ ^ distinct white 
r, radial vein; m, spot. Perhaps as severe a sufferer as 

medial vein; c cubital vein; a, anal ^ny other field product from the attacks 

veins. (After Comstock; enlarged.) ■' ^ 

of Noctuid larvae is cotton. The cotton- 
worm, Aletiaargillacea, feeds on the foliage of the cotton-plants and the cotton 
boll-worm, Heliothis armigera, attacks the cotton pods or bolls. These two 
caterpillars cause losses to the cotton-growing states of millions of doUars- 

FiG. 584. — Venation 
A gratis ypsilon. cs, 
subcostal vein 

^NA \i I ■ 'iiiML ■'!> 

. ^«»- ■!>.••• ^-*»4# v#"%i* ■-* ■*'n . 

Fig. 585. Fig. 586. 

Fig. 585. — Larvae of the gray dagger-moth, Acronycta occidentalis. (After Lugger;. 

natural size.) 
Fig. 586. — Gray dagger-moth, Acronycta occidentalis. (After Lugger; natural size.) 

every year. The cotton boll-worm is more or less familiar in states farther 
north, under the name of corn-worm, where it is found feeding on ears of 
green corn and on tomatoes. It is a naked, greenish-brown, dark -striped 
caterpillar. The moth has pale clay-yellow fore wings with a greenish tint, 
the hind wings paler. 

Among the most conspicuous of all the caterpillars are the not unfamiliar 
larvae of the tussock-moths, Lymantriidae, one common species infesting our 



i = Catocala parta. 
2=Basilona imperialis. 
3 = Apanresis virgo. 
4=Pseudohazis eglanterina. 
5 = Automeris io. 


Mary Welhnan^ del. 

The Moths and Butterflies 


Fig. 587. — The raspberry dagger- 
moth, Acronycta imprcssa. (After 
Lugger; natural size.) 

shade-trees in town and country and another, less common, attacking orchards 
and forest-trees. The caterpillars (Fig. 588) of Notolophus leucostigma , 
the white-marked tussock-moth, which is the shade-tree species, are about 
i\ inches long, very hairy, bright yellow with a blackish stripe along the back 
and one along each side, but chiefly conspicuous by a series of four cream- 
colored dense tufts of vertical hairs on the back, three long black hair pen- 
cils, two on the front part and one on the 
hind part of the body, and by the coral-red 
head and similarly colored two small pro- 
tuberances on the sixth and seventh abdom- 
inal segment which are scent-organs used 
to repel enemies. When full-grown these 
caterpillars pull the hairs from their body 
and mixing them with some silk make a 
grayish cocoon on the tree-trunks. The fe- 
male moth is wingless, light gray in color, 
and unusually long-legged for a moth; when issued she simply crawls out of the 
cocoon and lays her 300 to 500 eggs covered by a frothy-looking but firm sub- 
stance in a grayish mass on the outside of it. The males are ashy gray and 
have broad short wings, expanding i^ inches, the fore wings with darker wavy 
transverse bands, a small black spot near the tip, an obhque blackish stripe 
beyond it, and a minute white crescent near the outer hinder angle. The 

antennas are feathery, and the 
fore legs tufted with hairs. The 
best remedy for these pests is 
to gather the egg-masses in the 
winter and put them into a box 
with its top covered by mosquito- 
netting. In the spring the moths 
and the egg parasites which are 
numerous will hatch; the minute parasites will escape through the netting 
to go on with their good work, while the moths will be retained in the box 
and may be killed. 

The orchard and fruit-tree species, Parorgyia parallela, the parallel- 
lined tussock-moth, is winged in both sexes, the moths being dark gray with 
darker-colored wavy lines and spots. The caterpillars are gray with lon- 
gitudinal black stripes; short black tussocks are found on the back of seg- 
ments 4 to 7, a pair of long black pencils is at each end of the body, and on 
the back of each of segments 9 and 10 is a small pale-yellow scent-cup. 
The head is shining black. It feeds especially on plum-, crabapple-, and 

The most notorious member of the Noctuidae is the gypsy-moth, Ocneria 

Fig, 588. — Larva of the tussock-moth, Noto- 
lophus leucostigma. (After Felt, natural size.) 


The Moths and Butterflies 

dispar, a European species brought to Massachusetts in 1868, and from 
1890 to 1900 fought at the pubHc expense. A gentleman living in Med- 
ford, a town of Massachusetts, imported a number of different kinds of Euro- 
pean silk-spinning caterpillars in an attempt to find some species which 
might be bred in this country in place of the mulberry silkworm {Bombyx 
mori). Some of the moths escaped from his breeding-cages, and among 
them some gypsy-moths. In a very few years the species had increased to 
such numbers and spread throughout such an extent of woods that it seri- 
ously threatened the destruction of all the forest- and shade-trees in north- 
eastern Massachusetts. By 189 1 it was causing great injury to forest-trees 
over 200 square miles. So far it has been confined because of the whole- 
sale operations against it. The State has employed as many as 570 men 
at a time in spraying, egg-collecting, trunk-banding, etc., in the great fight 

Fig. 589. — The California oak-worm moth, Phryganidia californica. A, eggs on leaf; 
B, just-hatched larva; C, full-grown larva; D, pupa, or chrysahd; E, moth; F, Pimpla 
behrendsii, parasite of the larva. {B, much enlarged; D and F, twice natural size; 
others natural size.) 

against the pest and up to 1900 had expended over a million dollars in the 
struggle. The caterpillar when full grown is i^ inches long, creamy white, 
thickly sprinkled with black, with dorsal and lateral tufts of long black and 
yellowish hairs. The cocoon is very slight, merely a few silky threads. The 
male moths, expanding i| to 2 inches, are brownish yellow with smoky fore 
wings bearing darker irregular transverse lines and pale hind wings with 
darker outer margins. The females are large, expanding 2^ inches, and 
creamy white in color, with irregular transverse gray or blackish hnes. 

The Moths and Buttertiies 407 

In California is found a pretty pale-brownish moth that flutters weakly 
about the live-oak trees in early summer and late autumn, which has the 
distinction of being the only North American species in the family Dioptida:. 
The larvs of this moth feed chiefly on the leaves of the live-oaks and white 
oaks in the California valleys and the species may be called the live-oak 
moth, Phryganidia calijornica (Fig. 589). The moths expand about i inch 
and are uniformly pale brownish, with thinly scaled and hence almost trans- 
lucent wings. The male has a small yellowish-white ill-defined blotch on 
the center of each fore wing. The eggs are laid by the early summer brood 
of moths on the under side of the leaves of the oaks and the naked light- 
yellowish black-striped larvae feed until October ist on the tough leaves. 
Then they crawl down to the tree-trunks or to near-by fences or logs and 
change to a naked greenish-white or yellowish chrysalid with many black 
lines and blotches. The moths issue in from ten to twelve days after pupa- 
tion and lay their eggs again on the oak-leaves. But here is a curious fact. 
All the eggs laid on white-oak leaves by these autumn moths are doomed 
to death because just at the hatching-time the white-oak leaves fall and dry. 
The live-oak retains its leaves all winter and the larvae hatched on them 
feed and grow slowly through the winter, pupating in May and issuing as 
moths about June ist. Thus each year about one-fourth of the eggs laid 
by this species are wasted. The larvae from the eggs laid on the white oaks 
in the spring live because they have white-oak leaves all summer to feed 
on, but those of the fall brood which hatch on the white oaks all die. In 
some seasons this insect is so abundant as to defoliate the oak-trees in cer- 
tain localities twice during the year, but whenever the caterpillars get so 
numerous a certain small slender ichneumon-fly, Pimpla behrendsii, which 
lives parasitically on them becomes also very abundant (there being plenty 
of food for its young) and soon checks the increase of the moth. Out of 
144 chrysaHds of the moth which I once gathered but 11 moths issued, 
99 of the chrysalids giving forth ichneumon-flies and the rest dying from 
other causes. I have found the caterpiUar most abundant on the live-oaks 
{Q. agrijolia), but it occurs also on Q. lobala, Q. kelloggii, Q. diimosa, and 
Q. doiiglassi. 

A family represented in this country by only four species is the Peri- 
copidae. Three of these species are found only in the western states, the 
fourth in Florida. The single species of the four at all familiar to collectors 
is the beautiful and abundant Gnophala latipennis, with its two or three 
varieties. This moth expands about 2 inches and is black, with two 
large white blotches on the fore wing, each blotch subdivided by the black 
veins running through it and single large blotch on the hind wing. A 
variety common in California has the blotches smaller and pale yellowish. 

The wood-nymph moths, Agaristidae, of which about two dozen species 


The Moths and Butterflies 

are found in North America, include a few strikingly patterned moths not at 
all uncommon. The moth known as the eight-spotted forester, Alypia odo- 
maciilata (PI. VIII, Fig. 5; also Fig. 590), is common in the Atlantic states; 

Fig. 590. — Three eight-spotted forest-moths, Alypia 8-maculala, and one beautiful wood- 
nymph, Eudryas grata (the lowest). (After Lugger; natural size.) 

it expands about i^ inches, has deep blue-black wings, with two large sub- 
circular whitish-yellow spots on each wing, the spot nearest the base on 
the hind wing being much larger than the outer one. The patagia (shoulder- 
lappets) are often yellow and the legs marked with orange. The larvae, 

The Moths and Butterflies 409 

which are light brown with many fine black lines and one broad orange 
band across each segment and head and cervical shield deep orange with 
black dots, feed on the V^irginia creeper, sometimes on the grape, and often 
are so abundant as to injure the plants seriously. The caterpillar is nearly 
i^ inches long when full-grown, and burrows into soft or rotten wood to 
pupate, or failing this pupates on or just below the surface of the ground. 

The beautiful wood-nymph, Eudryas grata (Fig. 590) (classed by 
some entomologists with the Noctuidse), is very different in color and 
pattern, having milk-white fore wings broadly bordered and marked with 
brownish purple and with two indistinct brownish spots in the center. 
The under surface of these wings is reddish yellow. The hind wings are 
yellow with a pale purplish-brown border. The head is black and there 
is a wide black stripe along the back of the thorax, breaking up into a 
series of spots along the abdomen. The caterpillar is much like that of 
the eight-spotted forester and feeds on the same plants. "The moth, which 
is active at night and sometimes attracted to electric lights in large numbers, 
is very often discovered during the day upon the surface of the leaves of its 
food-plants. Its closed wings form a steep roof over its back, and its four 
legs, which have a curious muft'-like tuft of white hairs, are protruded and 
give the insect a very peculiar appearance." 

The grape-vine Epimenis, Psychomorpha epimenis, is a small velvety 
black Agaristid moth with a broad, irregularly lunate, white patch across 
the outer third of the fore wing and a somewhat larger and more regular 
patch of orange-red or brick-red on the hind wings. Its bluish caterpillar 
feeds on grape-leaves. 

Delicate and pretty are the little footman-moths, Lithosiidae, in their 
liveries of drab or slate, yellow or scarlet, and with their slender bodies 
and trimly narrow fore wings. The larvae of but few species are known; 
they mostly feed on lichens and have the body covered with short stiff 
hairs. Because these caterpillars are not injurious but little attention 
has been given to the life-history of the footman-moths, and the amateur 
has here an opportunity to add to our knowledge of insects in an order 
popularly supposed to be pretty well "worked out." 

The moths themselves although few in number of species are well dis- 
tributed over the country, although the southwestern and Pacific states 
have really more than their share. Two common eastern species are 
the striped footman, Hypoprepia miniata, and the painted footman, 
H. juscosa, each expanding about i inch. The first is brick-scarlet, with 
two longitudinal broad plumbeous bars and the distal half of a third on 
the fore wing and a broad outer slaty border on the hind wings. The 
latter has almost the same pattern, but the ground color is distinctly yellowish 
red in place of scarlet or brown-red. Another common eastern Lithosiid 

41 o The Moths and Butterflies 

is the pale footman, Crambidia pallida, expanding nearly i inch and 
drab all over; C. cephalica, found in Colorado and Arizona, expanding 
not quite an inch, has both wings and the whole body of a delicate shining 
silvery white. The banded footman, Cisthene (Ozonadia) unijascia, found 
all along the Atlantic and Gulf coasts, expands f inch and has the fore 
wings dark with a narrow curving yellow band and the hind wings with 
the base and disk pink or yellowish, the apex being dark. Lithosia (Lexis) 
hicolor, found in the northern states and Canada, expands nearly ij inches 
and is slate-colored, with yellow on the front margin of the fore wings, the 
tip of the abdomen, the prothorax, and the palpi. The several Rocky 
Mountain and desert species mostly have brick-red or drab or slaty ground 
color, some unmarked and some with dark border on the hind wings if 
red is the ground color, and smoky-whitish hind wings if body and fore 
wings are drab or slaty. 

Another family of moths expanding about an inch, and with a charac- 
teristic habitus due to the long narrow fore wings, the small size of the 
hind wings, and the contrasting colors of the wing-pattern, are the Zygasnidas, 
or SyntomidiE, as the newer nomenclature names them. In the hind wing, 
veins subcosta and radius are fused, usually for the whole length. About 
twenty species of the family are found in this country, and because, as 
with the Lithosiidce, the larvs are not of much economic importance the 
life-history of but few of the species is known. The majority of the species, 
besides, live in the western and southwestern states, and like other 
mountain, plain, and desert insects are hardly known except in their flying 
stage. The larvae of some species feed on grasses, of others on lichens. 

One of the most striking species is Cosmosoma auge, found in the 
extreme south, which has both fore and hind wings clear of scales over 
the base and disk only, a border all around the veins, and a small black 
patch at the tip of the discal cell of the fore wing covered with black scales. 
The plump body is scarlet, with the end of the abdomen and a dorsal 
longitudinal band on it metallic blue-black. The wings expand i inch. 
Lycomoipha is a genus of small Zyga^nids characterized by having the 
wings colored in two strongly contrasting shades, black and brick-red or 
black and reddish yellow. In L. pholus the basal two-fifths of each 
wing is yellow and all the rest black; in L. miniata the basal two- 
thirds is red, the rest black; in L. grotei all of the fore wing is red 
except a narrow black border on the outer margin, while the anterior 
half of the hind wings is red, the posterior half black. Ctenucha is a 
genus of larger species which have smoky-brown wings unmarked, as 
in C. virginica, a northeastern species, which has a yellow head 
and metallic bluish-black body, C. miiUijaria and C. ricberoscapus, 
Pacific coast species which have a coral-red head and shoulder-lappets 

The Moths and Butterflies 


and metallic deep-bluish body, or which have the fore wings marked 

by a few conspicuous longitudinal 

yellowish lines as in C. venosa, found 

in Colorado, New Mexico, and 

Texas. Scepsis julvicoUis, found in 

the eastern and Mississippi Valley 

states, has subtranslucent smoky 

wings with a region clear of scales 

in the middle of the hind wings; its 

prothoracic collar is yellow and its 

abdomen metallic blue-black. 

The "woolly-bear" caterpillars 
(Fig. 592) and the tiger-moths, which 
are the same insects in dilJerent 
growth stages, are among the most 
familiar of caterpillar and moth 
acquaintances. They belong to the 
family Arctiidae, represented in this 
country by a hundred and twenty 
species of which surprisingly many 
are pretty well known to any ardent collector. The strikingly colored, 
spotted, and banded wings of the stout and hairy-bodied moths and the 
dense clothing of long strongly colored hairs characteristic of most of the 

C a 

Fig. 591. — Venation of a Zygsenid, Ctenncha 
virginica. cs, costal vein; sc, subcostal 
vein; r, radial vein; ni, medial vein; c, 
cubital vein; a, anal veins. (After Com- 
stock; enlarged.) 

Fig. 592. — Woolly -bear caterpillars, Halesidota sp., all three of the same species but 
showing variations in extent of the black markings. 

larvae are the recognition-marks of the family. The moths, too, are 
mostly fairly large and are readily attracted by lights, while the cater- 

412 The Moths and Butterflies 

pillars, trusting to the uncomfortable mouthful of hairs they offer their 
bird enemies, travel conspicuously about in the open with a characteristic 
nervously hurrying gait. Thus the Arctians become familiar to collector 
and observer. 

The woolliest woolly bear is the larva, sometimes called "hedgehog," of the 
Isabella tiger-moth, Pyrrharctia (Isia) Isabella (PI. VII, Fig. 3), common all 
over the United States; it is covered with a stiff furry evenly shorn coat black at 
either end and red-brown in the middle, and is comnionly seen in the autumn 
traveling rapidly about in open places. It hibernates in larval stage under 
loose bark or logs or sidewalks, and, after a brief activity in the spring, pupates 
within a slight cocoon made up of silk and its own brown and black hairs. 
The moth which issues soon is dull orange with the front wings variegated 
with dusky and spotted with black; the hind wings are lighter and also 
black-spotted; it expands 2 inches. The caterpillars feed on various plants, 
sometimes becoming destructive, when in sufficient numbers, to black- 
berry and raspberry bushes and to nursery stock. Lugger says that they 
are especially susceptible to attack by muscardine, a parasitic fungus disease 
much feared by silkworm-growers. "Hedgehogs" killed by muscardine are 
found stiffly attached to their food-plants with a whitish powder over the 
body at the base of the dense hair covering. 

The yellow bears, common caterpillars on the leaves of vegetables, 
flowering plants, and fruits, distinguished by their dense but uneven coat 
of long creamy-yellow, light or even dark brown hairs, are the larvae of the 
beautiful snowy-white miller-moth, Spilosoma virginica. The wings bear a 
few (two to four) small black dots, and the abdomen is orange-colored 
with three rows of black spots. The larv« pupate in the fall in cocoons 
composed almost wholly of their own long barbed hairs, and the moths issue 
in the spring. There is usually a second brood each year. This moth is 
kept in check by many parasites, few other insects having to contend with 
so many of these insidious enemies of their own animal class. 

The most destructive member of the family is the fall web-worm, Hyphan- 
tria cunea, which makes the large unsightly silken "nests" in plum-trees, both 
wild and cultivated, so familiar in late summer and autumn. The eggs 
are deposited in regular clusters of 400 or more on the plum-leaves, and the 
hatching pale-yellow larvae spin small silky web-nests close together which 
finally get included in one large one. The full-grown larvae are pale yellow- 
ish or greenish with a broad dusky stripe along each side; they are covered 
with whitish hairs which rise from black and orange-yellow warts. They 
often hang from the nest or branches by a long silken thread. They pupate 
in crevices of the bark and other sheltered places on the ground, passing 
the winter in this stage. The milk-white moths, sometimes with small 
black spots on the wings, sometimes unspotted, issue in late spring or early 

The Moths and Butterflies 


summer. They expand ij inches. There is much variation in color and 
pattern in both moths and caterpillars, many varieties being found in a 
single tree. 

Among the most strikingly colored and patterned Arctians are the numerous 
species of Apanresis (Arctia). A. virgo (PI. VI, Fig. 3), a common species 
in the Atlantic states, whose larva feeds on pigweed and other uncultivated 
plants, expands 2^ inches, has black fore wings with the veins broadly marked 
with pinkish yellow, and red hind wings with large angularly irregular black 
blotches. The thorax is colored like the fore wings, the abdomen like the 
hind wings. Sharply angled black spots on a ground of reddish, pinkish, 
salmon, and yellowish characterize almost all the many species in this genus. 




k^ ^^Mi^^i^^M^ 
















i4j I (L* 






Fig. 593. — Caterpillar of Halesidota tesselata. (After Lugger; natural sj:c.) 

Striking moths are Arachnis pida (PI. VIII, Fig. 4), with whitish fore wings 
marked with wavy "band-like blotches of pearl-gray, and red hind wings with 
three uneven gray bands; Ecpantheria deflorata, the leopard-moth of the south 
Atlantic states, and E. muzina, of the southwestern states, both creamy white 
with circular or elliptical black spots or rings thickly scattered over the fore 
wings, but only in a single submarginal series on the hind wings; and Utethe- 
isa bella (PI. VII, Fig. 7), a familiar little moth of the Atlantic states with 


The Moths and Butterflies 

its pinkish-red hind wings with black branching border and yellowish-red 
fore wings crossed by six bending white bands containing small black spots. 
Attractive and familiar moths are the various species of Halesidota, whose 
larvae feed on the leaves of hickory, oak, and several kinds of orchard trees. 
These caterpillars (Fig. 593) are covered with short spreading tufts of hairs 
white and black or yellow, and bear, too, a single pair of long hair pencils 
usually black or orange. They are often called tussock-caterpillars and 
are not unlike the true tussock-moth larvae (see p. 404). The moths 

(Fig. 504) have long narrow fore 
wings, and hind wings only about 
half as fong; in H. tesseUata the 
hind wings are almost transparent 
yellowish (while the fore wings have 
faint darker short transverse lines 
or blotches) ; H. maculata (Fig. 595) 
has yellowish fore wings thickly 

Fig. 594. Fig. 595. 

Fig. 594. — Halesidota carya, above, and H. tesselala, below. (After Lugger; natural size.) 
Fig. 595. — Halesidota maculata. (After Lugger; natural size.) 

sprinkled with brown and blotched with creamy-white spots, the pale hind 
wings being unmarked; H. lobeniJa has the wings nearly transparent, the fore 
wings dusted with dark scales, and a regular check pattern on the front and 
hind margins, the hind wings unmarked, and the abdomen of a beautiful 
rose color; H. argentata has the fore wings blackish brown with distinct 
white spots all over the surface, white hind wings bearing a single irregular 
brown spot near the apex. The Callimorphas (Fig. 596) are pretty, slender- 
bodied Arctians with snow-white, creamy, or soft warm yellow-brown wings, 
banded with dark brown or blackish; they belong to the genus Hap'oa, 
whose larvae are blackish studded with blue spots, and covered with short 
stiff hairs. All the species of Haploa are found in the Atlantic states. H. 
clymene (PI. VII, F'g. 5) has the wings brownish yellow, paler on the fore wings, 
which are incompletely bordered with blackish brown, a curious blunt arm 
of this color projecting in from the hinder margin; the hind wings have a 
subcircular dark spot; H. lecontei has white hind wings, and brown fore 



i = Anisota rubicunda 
2=Geometra iridaria, 
3=Pyrrharctia Isabella. 
4=Tropoea luna. 
5 = Haploa clymene. 
6=Melittia ceto 
7=Utetheisa bella. 






Mary Wellinaii, del. 

The Moths and Butterflies 415 

wings with six large white blotches; H. julvicosta has all the wings pure 
white with the front margin of the fore wings weakly fulvous. A familiar 
Arctian is the salt-marsh-caterpillar moth Eustigme acrcea, expanse i^ 
inches, with creamy-white fore wings and soft yellow-brown hind wings, all 
the wings sparsely dotted with black. 

A small family which includes a few widely distributed and well-known 
moths is the Lasiocampida;, of which the tent-caterpillar moths are the most 
familiar. All the Lasiocampid moths, which are robust, hairy, and fairly 
large, lack the frenulum, having, however, the humeral angle of the hind 
wing expanded so as to overlap the inner hind angle of the fore wing. In 
this humeral angle are one or two short supporting veins or vein-spurs. 

Fig. 596. — Haploa fitlvicosta (above) and H. contigua (in the middle and below). 
(After Lugger; natural size.) 

The best-known eastern species is the apple-tree tent-caterpillar, the 
forest tent-caterpillar being also familiar; on the Pacific coast also occur 
two common species, one specially affecting orchard trees. These four 
species belong to the genus Clisiocampa (Figs. 598, 599); the moths expand 
about 1^ inches and are all brown, varying in shade from yellowish to walnut 
to chocolate-brown, with a pair of pale or distinct light or darker oblique lines 
on the fore wings. C. americana, the apple-tree tent-caterpillar, lays its 
three hundred eggs in the summer in a band or ring glued around a small 


The Moths and Butterflies 

twig of an apple or wild-cherry tree; the eggs do not hatch until the follow- 
ing spring, when the young larvae feed on the buds and young leaves of the 
tree. The social larvte build a little web or nest in the fork of a branch, 

going out of it only to feed. As the 
caterpillars grow they enlarge the web 
until it becomes a bulky ugly affair 
perhaps two feet long, partly filled with 
excrement and cast skins. The full- 
grown caterpillars are blackish with 
yellow and bluish spots, white striped 
along the back, and covered with fine 
yellowish hairs. "They feed on the 
young and tender leaves, and eating 
on an average two leaves a day the 
young of one pair of moths consume 
from ten to twelve hundred leaves, and 
Fig. 597.— Venation of Halesidota tessel- as it is not uncommon to find from six 

lata, cs, costal vein; sc, subcostal to eight nests on a single tree not less 

vein; r, radial vein; «^ medial vein; ^^^ seventy-five thousand leaves are 

c, cubital vein; a, anal veins. (Alter ■' 

Comstock; enlarged.) devoured, a loss which no tree can long 

endure." In about forty days the larvae 
are ready to pupate, when they scatter from the nest, find sheltered places 
under eaves, fence-rails, etc., and spin spindle-shaped cocoons of white, 
almost transparent silk, within which they change. After twenty to twenty- 
five days of pupal life the winged moths issue and soon after lay their 
eggs for next year's brood. The life-history of the various other species 
is similar to this although other trees are chosen for feeding-grounds. 

The lappet-moths, so-called from the curious lobes or lappets arranged 
along the sides of their caterpillars, are of several species. Tolype velleda, 
expanding 17 to if inches, has a white body with a black spot and dusky- 
gray wings crossed by white lines; its caterpillar feeds on the fohage of 
apple-, cherry-, and plum-trees, and is hair-fringed and protectively colored so 
that it looks much like an excrescence of the bark on which it habitually 
lies when not feeding. Gastropacha americana (Fig. 601), the American 
lappet-moth, expanding i| inches, is so hke a dead leaf in appearance that 
it can hardly be distinguished when at rest; it varies somewhat in color, 
but most individuals are reddish brown with a broad interrupted whitish 
band across both wings; the hinder and outer edges of the fore wings and 
the outer edges of the hind wings are deeply notched. The caterpillar feeds 
on apple, cherry, and oak, hiding during the day but becoming active at 
night. It is broad, convex above and flat beneath, ash-gray with fringes 
of blackish or gray hairs, and when at rest it is almost impossible to recognize. 

The Moths and Butterflies 


It grows to be 2 inches long and spins a peculiar gray cocoon which looks 
very much like a slight swelling of the twig to which it is fastened. The 
pupa hibernates, the moth issuing in June of the next year. 

Fig. 598. Fig. 599. 

Fig. 598. — A family of young forest tent-caterpillars, Clisiocampa disstria, resting during 
the day on the bark. (Photograph from life by Slingerland; one-third natural size.) 

Fig. 599. — The forest tent-caterpillar moth, Clisiocampa disstria, in its various stages. 
m, male moth; /, female moth; />, pupa; e, eggs in a ring about twig; g, eggs after 
hatching; c, larva or caterpillar. (After Slingerland; moths and caterpillar natural 
size, eggs and pupa slightly enlarged.) 

Including the largest, the most beautiful — in popular eyes at least — 
and the favorite moths for rearing in "crawleries," the superfamily Saturniina 
includes as well one of the only two insects that have been domesticated 
by man and reared for the sake of their useful products. The honey-bee 
and the silkworm moth are fairly to be called domesticated animals. To 
the Saturniina belong the great cecropias, the marvelous lunas, the regal 
and imperial walnut-moths, and the soft-tinted rosy dryocampas. Although 
the whole group, divided commonly into four families, includes but forty- 
two North American species, almost every one of these is more or less 


The Moths and Butterflies 

familiarly known to the amateur collector and crawlery owner. And popular 
books like Dickerson's "Moths and Butterflies," Eliot and Soule's "Cater- 
pillars and Their Moths," etc., which tell in 
detail of the life-history and habits of various 
Lepidoptera, mean by "moths," first Saturnians, 
then Sphingids, and finally a scant sprinkling 
of "others." The giant vividly colored cater- 
pillars, the great silken cocoons safely enclosing 
their mystery until that day when a marvel of 

a a 

Fig. 600. Fig. 601. 

Fig. 600. — Venation of Clisiocampa americana. cs, costal vein; sc, subcostal vein; 

r, radial vein; ;w, medial vein; c, cubital vein; a, anal veins. (After Comstock; 

Fig. 601. — The American lappet-moth, Gastropacha americana. (After Lugger; natural 


living color and pattern slowly crawls out and unfolds and takes on the 
seeming of the perfect cecropia or polyphemus, it is little wonder that the 
giant silkworm-moths are — always never overlooking the swift and masterful 
Sphingids — the moths of popular fancy. 

Just because these moths are so well known and so well and fully written 
of elsewhere I may limit my account of them to a brief descriptive catalogue 
of adults and larvae with the particular aim of making the more common 
species determinable by amateurs. The particular species in hand once 
safely identified, details of life-history and habits can be looked for in the 
many popular or technical accounts of the various kinds. In all, the males 
can be distinguished from the females by their large antennae and smaller 
bodies. In some species the sexes are very different in color and pattern. 

Of the genus Samia, the real giant silkworms, four species occur in 
this country. S. cecropia, the great cecropia-moth of the eastern states, 
expands 5 to 6 inches, has red thorax with white collar, red abdomen 
banded with white and black lines, wings with grizzled gray ground, and 
markings, as shown in Fig. 602, of reddish white and blackish with clay- 
colored outer margins. The large discal spots on the wings are whitish in 
the center, surrounded and encroached on by reddish, and margined with a 
narrow black line. The full-grown larva (Fig. 604) is nearly 4 inches 
long, pale limpid green, and bears on its back conspicuous tubercles, coral- 

The Moths and Butterflies 


red on the second and third thoracic segments, bkie on the first thoracic and 
last abdominal, and yellow on the others; smaller blue lateral tubercles are 
present. It feeds on many kinds of orchard- and forest-trees, most small fruits, 
and some herbaceous plants. The winter is passed in the pupal stage 
enclosed in a great pod-shaped rusty-gray or brownish silken cocoon about 
3 inches long and i inch wide in the middle, composed of two layers, 
an outer strong "brown-paper" layer and an inner loose fibrous one. The 
pupaj may be easily found on trees when the leaves are off and brought 

Fig. 602. — Cecropia-moth, Samia cecropia. (Photograph by author; natural size.) 

into the house. The moths will issue in early summer through an opening 
which is left by the larva in one end of the cocoon. S. Columbia of the north- 
eastern states and Canada is smaller than cecropia, the angulated discal 
wing-spots have hardly any reddish border and the transverse outer wing- 
border of white has no red outer margin as in cecropia, the abdomen is dark- 
red brown rather than red, and the basal half of the front wings is tinged 
with reddish brown. 5. gloveri, found in the Rocky Mountains and west 
to Arizona, is like Columbia, but as large as cecropia. S. ceanothi of the 
Pacific coast has the ground color of the wings strongly reddish, the outer 


The Moths and Butterflies 

markings weak to wanting, the white transverse wing-band narrow and 
with no reddish border, the discal spots also without reddish margin. 

The polyphemus-moth, Telea polyphe- 
miis (Fig. 605), expanse 4 to 5 inches, 
common in the whole country, is ocherous 
brown with a pinkish margined blackish 
outer transverse band across each wing 
and a discal spot on each wing with 
unsealed clear center; this latter char- 
acter makes the species at once unmistak- 
able; the hind wing-spots are in the center 
of a large blackish blotch with bluish 
scales by the inner margin of the clear 
spot. The larva (Fig. 606), which feeds 
on various forest-, shade-, and orchard- 
trees, reaches a length of 3 inches or 
more, is light green with seven oblique 
pale-yellowish lines on each side of the 
body, and bears numerous little black 
wart-like processes provided with small 
stiff bristles, and each body segment has 
a small silvery spot on the middle. The 
dense oval, completely closed cocoon is 
made of silk and a few leaves closely 
wrapped and tied together. It usually 
falls to the ground in autumn, but sometimes remains on the tree. The 
moth secretes a fluid from its mouth which softens and partly dissolves one 
end of the cocoon for its emergence. 

Fig. 603. — Venation of a Saturniid, 
Botnbyx mori. C5, costal vein; sc, 
subcostal vein; r, radial vein; m, 
medial vein; c, cubital vein; a, 
anal veins. (After Comstock; en- 

Fig. 604. — Larva of Samia cecropia. (After Dickerson; natural size.) 

In Plate VII, Fig. 4, is shown in color the luna-moth, or pale empress 
of the night, Tropcea luna (Fig. 607), a marvel of delicate green tinting 

The Moths and Butterflies 


and exquisite symmetry of curving outlines. It expands 4^^ inches, and 
ranges over the whole country. The larva is rather like that of the polyphe- 
mus-moth, being clear, pale bluish green with a pale-yellowish stripe on 

Fig. 605. — The polyphemus-moth, Telea polyphemus, and cocoon. 
(After Lugger; reduced about one-fourth.) 

each side of the body; each segment bears about six small purplish or rosy- 
tinged pearl tubercles; at the tip of the body are three brown spots edged 
with yellow. It feeds on hickory and walnut, on other forest-trees, and 

Fig. 606. — Larva of polyphemus-moth, Telea polyphemus. 
(After Dickerson; natural size.) 

makes a rather thin but compact cocoon of silk and leaves. 

In the eastern states the Asiatic ailanthus-worm moth, Philosamia 
cynthia, expanse 5 inches, with angulated wings, olive-brown ground-color 


The Moths and Butterflies 

on body and wings, a whitish lunate discal spot and a white and purplish 
transverse bar on each wing, and body with longitudinal series of white 
tufted spots, has become common near several cities. 

The promethea-moth, Callosamia pronielhea, expanse 3 to 4 inches, light 
reddish brown in female, and blackish and clay color in male, with mark- 
ings as shown in Fig. 609, is perhaps the most abundant of all these giant 
moths. Its larva when full-grown is 2 inches or more in length; it is bluish 
green and the body bears longitudinal series of black pohshed tubercles, 
two of these tubercles on each of the second and third thoracic segments 

Fig. 607. 

-The luna-moth, or pale empress of the night, Tropcea luna. 
(After Lugger; reduced about one-fourth.) 

being larger and red instead of black. It feeds on many kinds of trees, but 
Comstock has found it more frequently on ash and wild cherry than on 
others. The cocoon is long and slender and enclosed in a dead leaf whose 
petiole has been fastened to the branch with silk by the larva. "At the 
upper end of the cocoon there is a conical valve-like arrangement which 
allows the adult to emerge without the necessity of making a hole." C. 
angulijera is a moth slightly larger than promethea, but otherwise hardly 
distinguishable from it except that the shape and markings of the wings, 

The Moths and Butterflies 



Fig. 608. — Cocoons: i, 2, 3. of Tropma hma; 4, 5, 6, of Callosamia angtdifera; 7, 8, 9, lo^ 
oi Callosamia promethea. (After Laurent; somewhat reduced.) 


The Moths and Butterflies 

which vary a httle in male and female of promethea, are identical in this. It 
is found also only in the Atlantic states. 

The lo emperor-moth, Automeris io (PI. VI, Fig. 5; also Fig. 610), ex- 
panse 2^ to 3 inches, is the most familiar and the only eastern species of 
the four members of this genus. It can be recognized by the large blue 
and black eye-spots in hind wings and by its unmarked fore wings. The 
female has rich purplish-brown fore wings, the markedly smaller male yellow 
fore wings. The larva (Fig. 611), which feeds on trees, small fruits, corn, 
clover, etc., when full-grown is 2^ inches long, and is pale green with a 

Fig. 609. — The proinetliL-a-moth, Callusaiiiia promethea, male. 
(After Jordan and Kellogg; natural size.) 

broad brown stripe edged with white and reddish lilac on each side, and 
has the body covered with clusters of black-tipped green branching spiny 
hairs which are very sharp and strongly stinging. The thin, irregular 
parchment-like cocoon made of tough gummy brown silk is spun under 
dead leaves or other rubbish on the ground. In Texas is found A. zelleri, 
expanse 5 inches, reddish brown, without any yellow color in hind wings; 
in Arizona yl. pamina, expanse 2^ to 3 inches, with yellow around the white- 
centered black eye-spots of the hind wings; and in New Mexico ^. se/'/zyr/a, 
expanse 2^ to 3 inches, with brown-black fore wings and pale-brown abdomen 
broadly banded with red. 

With a single species, the maia moth, in the eastern states, and but half 
a dozen in the Rocky Mountains, desert and Pacific slope states, the genus 
Hemileuca presents a striking difference from the other Saturnians so far 

The Moths and Butterflies 


described in the thinly scaled, not hairy, condition of the wings and the 
prevalence of black and white in the pattern instead of warmer colors. H. 
maid, expanding 2^ inches, is subtransparent black with a broad middle 
transverse band of white on each wing; in this band is a small blackish blotch 

Fig. 610. — The lo emperor-moth, Automeris io, and cocoon; female moth above; 
male below. (After Lugger; natural size.) 

isolated in the hind wings, but connected with the black of the base in the 
fore wings. This species occurs in the eastern states; a similar species, H. 
nevadensis, being found from the Rocky Mountains to the Pacific; H. electra, 
found in southern California, has the hind wings blackish red; other species, 
found in New Mexico and Arizona, are mostly black and white with a red- 


The Moths and Butterflies 

dish or pinkish tinge here and there. The larva of H. maia feeds on oak; 
it is brownish black with a lateral yellow stripe, and has large branching 
spines over the body which sting severely. 

In Plate VI, Fig. 4, is shown in proper color and pattern a bizarre 
moth, Pseudohazis eglanterina, not uncommon in the Rocky Mountains, which 

Fig. 611. — Larva of lo emperor-moth, Automeris io. (After Dickerson; natural size.) 

we may call the clown. An allied species, P. sJiastaensis, similarly marked 
and colored, is found on the Pacific slope, and a third species, P. hera, with 
pale yellowish-white ground-color in the wings instead of purplish red, occurs 
in the region between the Rocky Mountains and the Sierra Nevada. 

Two great moths, the imperial (PI. VI, Fig. 2) and the regal walnut- 
moth (Fig. 612), are the most impressive of a subgroup of the Saturniina 
called the Ceratocampidae. They are all short-bodied and hairy and show 
for colors exclusively rich warm browns and soft yellows, light purple and 
rose. A curious structural characteristic of the family is the limiting of the 
pectinations on the antennae of the male to the basal half of the antenna. 
The regal walnut-moth, atheroma regalis (Fig. 612), expands fully 5 inches, 
has a rich brown ground-color on body and hind wings, with the fore wings 
slaty gray with yellow blotches, and veins broadly marked out in red-brown. 
The larva (Fig. 613), 4 to 5I inches long, and yellowish brown, reddish 
brown, or greenish, is distinguished from all other caterpillars by the great, 
threatening, but harmless blue-black horns of the body; it feeds on butter- 
nut, walnut, ash, pines, and other trees. Basilona imperialis, the imperial 
moth, is as large as the regal walnut, but with ground-color of rich yellow, 
overspread on base and outer part of fore wings and as a spot and band 
on hind wings with soft brownish purple. The larvae when full-grown are 
3 inches long, brown or greenish, thinly clothed with long whitish hairs, 
and bear conspicuous spiny horns on the second and third thoracic segments. 
They feed on hickory, oak, elm, maple, and other deciduous forest-trees, 
as well as on spruce, pine, juniper, and hemlock. The larvae of both these 

The Moths and Butterflies 


great moths burrow into the ground to pupate, the rough brown naked 
chrysalids wintering over. 

Fig. 612. — The regal walnut-moth, Citheronia regalis. (Photpgraph by author; natural size.) 

Anisota is a genus of smaller moths containing five species limited to 
the eastern states, four of which are brown and one, A. riibiciinda, rosy and 

Fig. 613. — Larva of regal walnut-moth, Citheronia rcgaus. 
(Photograph by author; natural size.) 

yellow. This latter, called the rosy dryocampa, is shown in color in Plate 
VII, Fig. I. Its larva, so:r:^t!n^es ca.led the green-striped maple-worm, 


The Moths and Butterflies 

is pale yellowish green and is striped with many fine longitudinal lines 
alternating lighter and darker than the ground-color. There are two horns 
on the second thoracic segment, and dorsal spines on the eighth and ninth 
abdominal segments. 

A. virginiensis is purpHsh red or brown, and the wings are nearly trans- 

FiG. 614. Fig. 615. 

Fig. 614. — The orange-striped oak-worm moth, Anisota senatoria, male. (After Lugger; 

natural size.) 
Fig. 615. — The orange-striped oak-worm moth, Anisota senatoria, female. (After 

Lugger; natural size.) 

parent in the center; the larva, found on oak, is grayish or greenish with 
brownish-yellow or rosy stripes and with small white warty processes all over 

Fig. 616. — Mulberry silkworms, larvae of Bombyx mori. (From life; natural size.) 

the skin; A. stigma, expanse 2 inches, is light ocherous brown with many 
blackish dots; its bright tawny or orange caterpillar has long spines on 

The Moths and Butterflies 


the back; A. senatoria (Figs. 614 and 615) is like /I. virginiensis, but lacks 
the transparent place in the middle of the wing; the caterpillar is black with 
four stripes. All these Anisota larvai feed on oaks, and that of A . senatoria 
also on blackberries and raspberries. Sphingicampa (Adelocephala) hicolor 
is a beautiful moth with brown fore wings and dark-pink hind wings with 
dusky dots, which is not uncommon in the Mississippi Valley and southern 
states; its larvae feed on the locusts and the Kentucky coffee-bean. In the 
southwest are two or three species of the genus Syssphinx resembling Sphingi- 
campa bicolor, but one, S. heiligbrodti, in Arizona, has iron-gray fore wings. 
Now unknown in wild condition, the long-cultivated Chinese or mulberry 
silkworm, Bombyx mori, is spread over most of the world, living exclusively, 
however, under the personal care of man. Indeed it is often said that the 
worm is so degenerate, so susceptible to unfavorable circumstances, that 
it could not live out of doors uncared for. As a matter of fact, however, I 
have bred moths from silkworms placed 
exposed on mulberry-trees in California 
immediately after the first moult. And 
these individuals experienced consider- 
able hardship in the way of low temper- 
atures and dashing rains. The heavy 
creamy-white moths, with wing expanse 
of if inches, take no food at all, and 
most of them cannot even fly despite 
their possession of well-developed wings, 
so degenerate are the flight-muscles from 
generations of disuse. The eggs, about 300, are laid by the female on any 
bit of cloth or paper provided her by the silkworm-growers. They are yellow 
at first, but soon change to a slaty color due to the beginning development 
of the embryo. In the annual race of silkworms, i.e., the variety which 
produces but one generation a year as compared with those others which 
produce two (bivoltins), three (trivoltins), and even five or six (multivoltins), 
the development of the eggs soon ceases, and they go over the winter, hatching 
in the following spring at the time the mulberry-trees begin leafing out. 
The larvae (Figs. 616 and 617) must be well fed with fresh mulberry or osage- 
orange leaves (they may at a pinch be carried through on lettuce) from which 
all rain- or dew-drops should be wiped off. The worms moult every nine 
or ten days, ceasing to feed for a day before each moulting, during the forty- 
five days of larval life, spinning before the last moult (pupation) the dense 
white or golden silken cocoon which is, to man, the silkworm's raison d'etre. 
In this spinning the thread is at first attached irregularly to near-by objects, 
but after a sort of loose net or web has been made the spinning becomes 
more regular, and by the end of three days a thick firm symmetrical closed 

Fig. 617. — Mulberry silkworm, show- 
ing front view of head and thorax. 
(From life; natural size.) 


The Moths and Butterflies 

cocoon, composed of a single continuous silken thread averaging over looo feet 
long, is completed. Inside this cocoon the larva pupates, and if undisturbed 
the chrysalid gives up its damp and crumpled moth after from twelve to fourteen 


days or longer. A fluid secreted by the moth softens one end of the cocoon 
so that the dehcate creature can force its vi^ay out. But this is not the usual 
fate of a silkvirorm pupa. The professional. grower must save the cocoon 

The Moths and Butterflies 


from injury by the moth, so he kills his thousands of pupae by dropping 
the cocoons into boiling water or by putting them into a hot oven. Then, 
after cleaning away the loose fluffy silk of the outside, he finds the beginning 
of the long thread which makes the cocoon, and with a clever little reeling- 
machine he unwinds, unbroken, its hundreds of feet of merchantable silk floss. 
From here to the silk-dress stage is a story not entomological, but one of 
elaborate machines and processes of human devising. 

Hovering, humming-bird-like, in the early dusk over the deep flower- 
cup of a petunia or honeysuckle or great jimson-weed, with its long flexible 
proboscis thrust deep down to the nectaries, and the swift wings making a 

Fig. 619. — Larva of the achemon sphinx-moth, Philampelus achemon. 
(After Lugger; natural size.) 

faint haze on either side of the trim body, the sphinx-moth, or hawk-moth, 
or humming-bird moth, as variously called, is a familiar garden acquaintance. 
But that he is but one of a hundred different American species; that he has 
cousins red and cousins green, somber cousins and harlequin cousins; that, 
strong-winged, clean-bodied, exquisitely painted, and honey-fine in his taste 
as he is now, his earliest youth was passed as a "disgusting," soft, fat, green 
tomato-worm or tobacco-worm or grape-vine dresser, and that at a later 
adolescent period he lay buried in the ground, cased, mummy-like, in a dark- 
brown sarcophagus — all this may not be as familiar. Still, excepting the 
giant silkworm-moths, the Saturnians, no other moth group is so much 
affected by collectors and crawlery proprietors as the Sphingida?. Thus 
the various adolescent stages of several hawk-moth species are known to 


The Moths and Butterflies 

many amateurs, and numerous different sphingid species will be found in 
any collection of Lepidoptera. The uniformity of structural character in 
larvae and adults of the various species, and the general similarity of habits 
and life-history, make the family a coherent one, and one readily distinguish- 
able from other moths. These moths, with few exceptions, have long, nar- 

FiG. 620. — Larva of the sphinx-moth, Phlegethontitis Carolina. (After Jordan and 
Kellogg; one-half natural size.) 

row, pointed fore wings, very small hind wings, a smooth-coated, compact, 
cleanly tapering body, and a long proboscis, coiled when not in use, like 
a watch-spring, on the front of the head (Fig. 509). The colors and pat- 
terns are extremely varied, but uniformly quietly beautiful and harmonious. 

Fig. 621. — Larva of Phlegethontius celeiis. (After Soule; somevsrhat reduced.) 

The larvae (Fig. 619) are naked, usually green, often with repeated oblique 
whitish lines on the sides, and bear a conspicuous sharp-pointed horn, 
or, in fewer instances, a fiattish, button-like shining tubercle, on the back 
of the eighth abdominal segment. The caterpillars, or "worms," feed on 

The Moths and Butterflies 


the foliage of various plants, and when full-grown most of them descend 
and burrow into the ground to pupate. The chrysalid is naked, with firm, 
dark^brown wall, and is distinguished by the odd jug-handle-Hke sheath 
for the developing long imaginal proboscis. A few larvae pupate on the 

Fig. 622. — Pholus achenion, above, and Pholus pandorus, below. 
(After Lugger; natural size.) 

ground in a slight cocoon made of silk and a few leaves tied together. The 
insects hibernate in the pupal stage; a few are said to be double-brooded. 
The name sphinx, applied to these moths by Linnaeus a century and a half 
ago, is suggested by the curious attitude assumed by the larvae when dis- 
turbed; the front part of the body is lifted (Fig. 620) clear of the object 
on which the insect is resting, and the head is bent forward on the thoracic 
feet. This position may be held rigidly for hours. 

Of the many species found in this country we can refer to but a few of 
the more familiar or beautiful or interesting ones, and these references may 
be made brief because of the colored figures which are grouped in our frontis- 
piece. These figures render descriptions unnecessary. 


The Moths and Butterflies 

Best known of all the hawk-moths, both in larval and adult stage, are 
the five-spotted sphinges, the tomato- and tobacco-worm moths, Phlege- 
ihontius quinquemaculata (celeus) and P. sexta {Carolina) (PI. VIII, Fig. 3). 

Fig. 623. — Larva of Pholus achemon. (After Soule; natural size.) 

Quinquemaculata is the commoner in the north, sexta in the south; in both 
the larva (Figs. 620 and 621) is green with oblique white stripes on the side 
and a long sharp caudal horn, and feeds on tomato-, tobacco-, and potato- 
leaves or jimson-weed. The horn of sexta is red, 
that of quinquemaculata green or blue-black. 
The pupae are long and slender and dark 
brown (green at first), and are often found when 
plowing or digging up fields in which these plants 
have been grown. The moth of P. quinquemaculata 
has ashy-gray wings, with zigzag markings, while 
the wings of sexta are not thus marked. The 
great pandorus sphinx, Pholus (Philampelus) 
pandonis (PI. I, Fig. i), found in the eastern 
and central states, is one of the most beautiful 
of all moths. The larvas feed on grape-vines 
and Virginia creeper, and, measuring four inches 
long when full-grown, are rich reddish brown 
with five conspicuous cream-colored spots along 
Fig. 624. — Grape - vine each side; a shining black eye-like tubercle takes 
spliinx - moth, Ampelo- ^j^g j^^^ ^f ^ caudal horn. It pupates under- 
phaga myron. (Natural ^ /tt- < \ vu 1 • 

size.) ground. P. achemon (rig. 022), with markings 

much like pandorus, but with strong rosy color- 
ation instead of greenish, has a larva which also feeds on grape and Vir- 
gina creeper and may be recognized by its six (instead of five) lateral 
cream-colored blotches. 

JTIV .-1 



i = Deilephila lineata. 
2=Chser()campa tersa. 
3 = Phlegethontius sexta. 
4=Arachnis picta. 
5 = Alypia octomaculata. 
6=Anato]mis grotci. 
7 = Plusia simplex. 


niarv Welhnan^ del. 

The Moths and Butterflies 


The beautiful little Ampdophaga myron, with soft red-brown hind wings 
and brownish-gray fore wings, patterned as shown in Fig. 624, has a pea- 
green, cream-banded, and yellow and lilac spotted larva known as the hog- 
caterpillar of the vine, so named from its form — the third and fourth seg- 
ments being greatly swollen, the head and first two segments small— and 
its destructiveness to grape-vines. When ready to pupate it spins a brown 
silken open-meshed cocoon on the ground under leaves or other rubbish. 

Fig. 625. — The double-eyed sphinx, Smerinthus geminatiis, above; Paonias excacatus, 
in middle; and P. myops, below. (After Lugger; natural size.) 

A. versicolor (PI. I, Fig. 3) is a beautiful cousin of myro7i with greenish 
overlaid on the brown. An extremely slim, slender-bodied, and slender- 
winged sphinx is Chcerocampa (Theretra) tersa (PI. VIII, Fig. 2), found in the 
northern states. It is very swift. An abundant and familiar hawk-moth found 
all over the United States is the white-lined sphinx, Deilephilalineata (PL VIII, 
Fig. i). Its caterpillar feeds on various plants, as grape, apple, watermelon, 
buckwheat, turnip, and purslane; the latter seems to be the preferred plant. 


The Moths and Butterfiies 

Exceedingly variable in color and pattern, it is usually yellow-green with a 
conspicuous longitudinal row of elliptical spots on each side of the back, 

Fig. 626. — Larva of Smerinthus geminatus. (After Lugger; natural size.) 

each spot consisting of two curved black lines enclosing a bright crimson 
blotch and a pale-yellow line; all the spots are connected by a pale-yellow 

P'iG. ()27. — Sphinx gordius. (After Lugger; natural ^izc.) 

line edged above with black. Sometimes the larvae are black, with a 
narrow yellow line along the back and a series of paler- and darker-yellow 

The Moths and Butterflies 


spots. The double-eyed sphinx, Smerinthus geminatiis (PI. I, Fig. 2; also 
Fig. 625), is a common species whose larvae feed on apple, plum, ash, willow, 
birch, and other trees; the full-grown caterpillar (Fig. 626) is 2\ inches 
long, apple-green, with seven oblique yellow stripes on each side of the 
body and a violet caudal horn. The genus Sphinx (Fig. 627) contains 
nearly twenty species, all of them soberly patterned with grayish, brownish, 
and blackish, and most of them expanding more than three inches. 

Fig. 628. — Larva of the abbott-sphinx, Thyreus abbotti. {Alitr Soule; natural size.) 

While most hawk-moths have narrow tapering fore wings and a slender 
tapering smooth-coated body, structural conditions indicating a well-de- 
veloped flight power, a familiar species, the modest sphinx, Marumba modesta 
(PI. I, Fig. 4), found all over the country, is hairy, heavy-bodied, and 

Fig. 629. — Larva of abbott-sphinx, Thyreus abbotti; note difference in pattern from 
larva shown in Fig. 628. (After Soule; natural size.) 

broad-winged. The full-grown larvae are 3 inches and more long, whitish, 
yellowish, and bluish green, with fine white dots all over the skin; the cau- 
dal horn is short. They feed on "balm-of-Gilead," poplar, and other trees. 
Another species of unusual shape is the beautiful dark-brown and canary- 
yellow small tufted-bodied abbott-sphinx, Thyreus (Sphecodina) abbotti 
(PI. I, Fig. 6), found in the Atlantic and Mississippi Valley states. Its 
larvae (Figs. 628 and 629) feed on woodbine and grape. They are ''ashes- 
•of-rose" color, finely transversely lined with dark brown and with longitu- 
dinal series of brown blotches. They have a large circular, eye-like tubercle 
in place of a caudal horn. They may appear in two different patterns as 


The Moths and Butterflies 

shown in Figs. 628 and 629. The pupa is found under dead leaves or other 
rubbish. Very similar in appearance and habits is the grape-vine amphion, 
Amphion nessiis (Fig. 630), of the same size and shape and colors and found 

Fig. 630. Fig. 631. 

Fig. 630. — The grape-vine amphion, Amphion nessus. (After Beutenmiiller; natural 

size, i|-2 inches expanse of wings.) 
Fig. 631. — Larva of clear-winged sphinx, Hemaris difflnis. (After Soule; natural size.) 

in the same states; it may be distinguished, however, by a pair of conspicu- 
ous narrow, bright-yellow bands across the abdomen. The larvae are pale 
yellowish green or chocolate-brown with various obscure darkish stripes. 

Fig. 632. — The death's-head sphinx-moth; note skull-like markings on thorax between 
wings. This moth is looked on with superstitious dread by many people. (Photo- 
graph by author; natural size.) 

A few sphinx-moths have the wings partly clear. These are called the 
clear-winged sphinxes and belong to the genus Hemaris. H. thysbc (PI. I, 

The Moths and Butterflies 


Fig. 5) is the most abundant Eastern species, although H. diffinis, with 
bright-yellow hairs in place of brownish yellow on thorax and abdomen, is 
common. In Colorado and Utah is found a smaller species, H. brucei, 
with yellowish thorax and abdominal band, and in California are one or two 
varieties of H. diffinis. The larva of H. diffinis (Fig. 631) feeds on honey- 
suckle and snowberry-bush and is pale green above, darker green on the 
sides, with three brown stripes on the under side; the caudal horn is yellow 
with blue-black tip; some of the caterpillars, as is common among the larvae 
of this family, are brown instead of green. It is two-brooded. Moths just 
issued from the chrysalid have scales over all of the wing surface, but these 
scales are so loosely attached on the discal area that the first few flights 
dislodge them, so that the "clear- wing" comes about. The larvae of 
H. thy she feed on viburnum, snowberry, and hawthorn. 


Taken all in all the butterflies are the most familiar and attractive insects 
to people in general; their size, beautiful color-patterns, and daytime flight 

Fig. 633. — The Parnassian butterfly, Parnassius smintheus, which lives in the Rocky 
Mountains and Sierra Nevada at an altitude of 5000 feet and more. (Natural size.) 

chiefly account for this. Six hundred and fifty butterfly species (compare 
with the six thousand species of moths) are accredited to this country in 
the latest authoritative catalogue of North American Lepidoptera. These 
represent, according to this catalogue, thirteen families; a more usual classi- 
fication, however, groups all these species into six families. As this latter 
arrangement is in use in most of the insect manuals, it will be adopted in this. 
Comstock, who has given the classification of the Lepidoptera much attention, 
gives the following key to families: 


The Moths and Butterflies 

Fig. 638. Fig. 637. 

Fig. 634. — Venation of a Hesperid, Epargyreus tityrus. (After Comstock; enlarged.) 
Fig. 635. — Venation of a Papilionid, Papilio polyxenes. (After Comstock.) 
Fig. 636. — \ena.\.\onoi s.'HymphsiMd, Basilarchia astyanax. (After Comstock; enlarged.) 
Fig. 637. — Venation of a Lycaenid, Chrysophaniis thoe. (After Comstock; enlarged.) 
Fig. 63S. — ^Venation of a Pierid, Pontia protodice. (After Comstock; enlarged.) 
For all: cs, costal vein; sc, subcostal vein; r, radial vein; m, medial vein; c, cubital 
vein; a, anal veins. 

The Moths and Butterflies 441 


A. With the radius of the fore wings five-branched and with all of these branches 
arising from the discal cell (Fig. 634); club of antennae usually terminated by a 

recurved hook (Skippers.) Supcrfamily Hesperiina. 

B. Head of moderate size; club of antennas large, neither drawn out at the tip 
nor recurved. Large skippers with wing expanse of 2 inches or more. 

Megathymid^ (p. 441). 

BB. Head very large; club of antennce usually drawn out at the tip and with a 

distinct recurved apical crook. If the crook is wanting, the species expand 

less than 1} inches Hesperiid/E (p. 442). 

AA. With some of the branches of radius of the fore wings coalesced beyond the apex 
of the discal cell (Fig. 635); club of antennae not terminated by a recurved hook. 

(The butterflies.) Superfamily Papilionina. 
B. Cubital vein of the fore wings apparently four-branched (Fig. 635); most of 
the species with tails on the hind wings. 

(The swallow-tails and parnassians.) Papilionid.e (p. 446). 
BB. Cubital vein of fore wings apparently three-branched (Fig. 636). 

C. With only four well-developed legs, the fore legs being unused, much 

shorter than the others, and folded on the breast like a tippet, except 

in the female of Hypatus; radius of fore wings five-branched (Fig. 636), 

(The brush-footed butterflies.) Nymphalid^ (p. 450). 

CC. With six well-developed legs; radius of fore wings, with rare exceptions, 

only three- or four-branched (Fig. 637). 

D. Medial vein of the fore wings arising at or near the apex of the 
discal cell (Fig. 637), except in Feniseca tarquinius, in which the 
wings are dark brown with a large fulvous spot on each. 

(The blues and coppers.) Lyc^nid^ (p. 443). 
DD. Medial vein of the for^, wings united with last branch of radius 
for a considerable distance beyond the apex of the discal cell (Fig. 
638); ground color white, yellow, or orange. 

(The whites and sulphurs.) Pierid^ (p. 444). 

The family Megathymidse, or giant-skippers, contains but one genus, 
Megathyma, represented by but five species, of which none is found outside 
of the southern and southwestern states. The best-known and most widely 
distributed species is the yucca-borer, M. yucccr, whose larvae live as bur- 
rowers in the roots of several species of yucca, and are from 4 to 6 inches 
long. The eggs are laid on the leaves and the young larva? spend a short 
time above ground in a cylinder made of a rolled leaf tied across with silk. 
Later they tunnel into the stem and downwards into the root, sometimes to 
a distance of 2 feet or more. When ready to pupate they crawl up to 
the chimney-like funnel at the top of the burrow and transform there. The 
moth expands 2^ inches, is deep umber-brown with a notched ferruginous 
band and other smaller blotches on the fore wings, and the hind wings with 
a ferruginous border. The other giant-skippers are of similar size and 

442 The Moths and Butterflies 

markings, and all of them are more moth-like than butterfly-like in general 
appearance. They may be looked on, indeed, as a sort of connecting link 
between the moths and the true butterflies. 

The Hesperidae, or skipper-butterflies (PI. IX), are a great family of small^ 
big-headed, robust-bodied butterflies of obscure patterning in browns and 
blackish (a few forms white and dark gray). Nearly two hundred species 
are known in this country, but few of them are at all familiarly recognized 
as distinct species; general collectors and amateurs know them better 
grouped into generic units, as Erynnis, Amblyscirtes, Eudamus, Thorybes, 
Pholisora, etc. Indeed, but few professional entomologists feel competent 
to undertake the identification of Hesperid species. A few weU-marked or 
specially numerous and wide-spread forms are, however, fairly well known. 
The caterpillars of all have large heads, constricted necks, and bodies thick 
in the middle and tapering both ways, and often make protecting nests of 
leaves and silk. The silver-spotted skipper, Epargyretis titynis (PI. V, 
Fig. 3), is abundant over all the country and is readily recognizable by 
its large size and distinctive pattern; the broad, irregular, silver spot is on 
the under side of the hind wing. The caterpillar feeds on various Legu- 
minosae, especially wistaria and locust, and when full-grown is i| inches 
long, with large, ferruginous head bearing two large orange spots, and lemon- 
green body transversely banded with darker green; it builds a nest or case 
of leaves, in which it remains when not feeding; it pupates either in this 
larval nest or makes a loose cocoon somewhere on the ground, hibernating 
in this stage. Another of the larger species is the cuiious long- tailed skipper, 
Eudamus proteus, found in the south Atlantic states (ranging as far north 
as New York City) and distinguished by the tailed hind wings and iridescent 
green-brown color. The genus Hesperia includes a dozen or more specieS' 
which are thickly white-spotted on a blackish-brown ground, giving them 
a checkered gray appearance; most of these checkered skippers are limited 
to the western states, but one, H. tessellaia, is found commonly all over 
the country. It expands i^ inches, and has even more white than dark on 
the wings; it flies rapidly about close to the ground and lays its eggs on 
various mallows; the larva is green with a dark interrupted dorsal line, dark 
lateral bands, and a pale band below the spiracles. 

A whole host of skippers are the " sooty -wings," members of several 
genera, but almost impossible to be distinguished by means of written 
descriptions. They vary in size from an expanse of i inch to nearly 2 inches, 
and have the wings grayish brown to blackish brown to truly sooty, usually 
with obscure indications of markings on both wings and almost always 
with a few small distinct white spots near the apex of the fore wings. The 
small sooty-wing, Pholisora catidlus, common in the east, expands i inch 
and has uniformly nearly black wings with a few distinct white dots on 


Fig. I 













Painphila lioboniok, male, upper side. 
" " under side. 

" " female, under side. 

" upper side. 
' ' zabulon, male, upper side. 
" " " under side. 

" " female, under side. 

" " " upper side. 

" scudderi, male, upper side (type). 
" " female, upper side (type). 

" bcUus, male, up{)er side. 
" underside. 
" panoquin, male, upper side. 
" underside. 
" stigma, male, upper side (co-type). 

' ' pittacus, male, upper side. 
" " under side. 

' ' rhesus, male, upper side. 
" " " underside. 

' ' nemorum, male, upper side. 
" massasoit var. suffusa, male, under side. 
' ' draco, female, under side. 
' ' loammi, male, under side. 
" alcina, male, upper side (type). 
' ' panoquinoides, male, upper side (type) 
TEgiale .streckeri, male, upper side. 
Archonias lyceas, upper side. 


The Moths and Butterflies 


the fore wings. Several large species, known as dusty-wings, expanding 
i^ to if inches, with grayish-brown to blackish-brown wings, belonging to 
the genus Thanaos, are common. Another large group of nearly indis- 
tinguishable species is that of the Pamphilas (PI. IX). These skippers are 
mostly tawny and are specially recognizable by a discal black patch in male 
specimens, which appears like an oblique scorched streak near the center 
of each fore wing. This patch contains certain peculiar scales which give 
off scent presumably attractive to the females. Erynnis sassacus (PI. X, 
Fig. 5), common in the Atlantic states, is a good example of the group. 
The least skipper, Ancyloxypha numitor (PI. V, Fig. 5), is the smallest 
commonly seen and differs from other skippers in lacking the recurved 
hook at the tip of the antennae and in having a slender body. The 
pale-yellow pilose larva feeds on grasses, especially those that grow in wet 

The small butterflies popularly known as blues, coppers, and hair- 
streaks compose the family of Lyca?nidai, or gossamer-winged butterflies, of 
which a hundred and twenty-five species are recorded from the United States, 
mostly the western half. The popular names express well the colors and 
pattern characteristic of the group. They are delicate, light-winged, slender- 
bodied butterflies rarely expanding more than an inch and a half and either 
bluish (pale whitish blue to brilliant metaUic dark blue) or coppery or reddish 
or dark brown, often with small blackish spots, or marked with short fine 
little lines, hair-streaks, on the under side of the wings, and often with dehcate 
little tail-like processes projecting from the hinder margin of the hind wings. 
The larvae are flattened, short, broad, small, forked, slug-like caterpillars 
with small retractile heads; those of a few species distinguish themselves 
from all other butterfly larvae by feeding on other insects, especially aphids. 
The chrysalid is naked, suspended from the posterior tip and supported by 
a silken line, or "bridle," about its middle. 

Often to be seen fluttering or clustered about wet spots in the roadway 
are numbers of delicate little pale-blue butterflies with under side of w^gs 
almost white and conspicuously dotted with small black spots and with 
white-ringed slender antennae; these are "blues," some species of the old 
genus Lycaena now broken up by modern systematists into a half dozen or 
more different genera. The spring azure, Cyaniris pseudargiolus (PL V, 
Fig. 4), is a wide-spread and common example of the group; with its several 
varieties it ranges over the whole continent, and it is one of the few "blues" 
whose young stages are known. The larvas, which curiously secrete honey- 
dew from little openings on the seventh and eighth abdominal segments, feed 
on the "buds and flowers of various plants, especially those of dogwood 
(Comics), Cimifuga, and Actinomeris." As many as three broods appear 
in a year. The various species of blues differ slightly in size, in shade of 

444 The Moths and Butterflies 

coloring, as grayish blue, lilac-blue, purple-blue, etc., in number and distinct- 
ness of the small black spots, but only an expert can determine the 

Less in number of species and perhaps not quite so familiar are the 
"coppers" with orange, red-brown or dark-brown wings conspicuously 
spotted with black. Fig. 4 of PI. X shows the color, markings, and size 
of a typical "copper," Heodes hypophlaas, "one of the commonest butter- 
flies in the United States." Most of the other coppers have, however, hardly 
as bright-red a ground color on the fore wings, some being really somber. 
Most of them, too, are a little larger than hypophlceas. A species patterned 
and colored much like hypophlceas, but a half larger, is Chrysophanns thoe, 
found in the Atlantic states and west to the Rocky Mountains. The har- 
vester, Feniseca tarquinius, small, with bright orange-yellow above spotted 
with black and mottled gray and brown underneath, is a common species 
all through the eastern states west to the Mississippi River; its larva feeds 
on the woolly plant-lice like the alder blight, apple-tree aphid, etc. 

The hair-streaks, mostly belonging to the genus Thecla, have short narrow 
lines or streaks on the under sides of the wings, and are usually provided 
with one or more delicate little "tails" on the hind wings. They vary in 
color from a dull brown to a splendid glancing blue or blue-green. They 
usually have one or more reddish spots at the base of the "tails" and the 
under sides of the hind wings are often greenish or parti-colored. Thecla 
halesus, the "great purple hair-streak" (PI. V, Fig. 9), is our largest 
species, and is found in the southern half of the country. Like the blues 
the hair-streaks are very difiicult to classify to species; indeed professional 
entomologists are not at all satisfied with our present systematic knowledge 
of the Lycaenids. 

In the extreme southwest are found rather rarely the few species of 
"metal-marks," Lemonias and Calephelis, black and reddish checkered 
Lycaenids, which occur in this country. Sometimes, as in L. virgiilti, the 
wings are spotted with white. The vernacular name is derived from a few 
small lead-colored or pearly-white spots near the outer margin of the wings. 
The tiny metal-mark, Calephilis ccBnius, expanding only | inch, and with 
the reddish-brown wings spotted with small steely-blue markings, comes 
as far north as Virginia. 

A smaller family than the Hesperida? or Lyca?nida?, but with numerous 
better-known members, is the Pieridae, the whites, yellows, and orange- 
tips. Because the larvae of several species feed on cabbage and other 
cruciferous plants, the unhappy name of cabbage-butterflies is sometimes 
applied to them. The common whites and yellows are the most famihar 
of roadside butterflies, but of the sixty species composing the family in this 
country, only half a dozen occur in the northeastern states, the south and 

The Moths and Butterflies 445 

west being the favored regions of distribution. All the species except two or 
three are of medium size, that is, have an expanse of ij to 2 inches, and 
have white or yellow, from light sulphur to orange, as ground color, with 
markings of black. The larva) are mostly green, longitudinally striped, 
with more or less distinct lines usually paler, and harmonize so thoroughly 
in coloration and appearance with the green foliage on which they feed that , 
they are not often seen. The chrysalids are naked, supported at the pos- 
terior tip and also by a loose silken bridle, and distinguished from other 
butterfly pupai by a conspicuous median-pointed process on the head end. 
The males of many Pierids give off a pleasing aromatic odor which comes 
from certain scent-scales (androconia) scattered about over the wing-surface. 
If the fore wings of a freshly caught male cabbage-butterfly be rubbed 
between thumb and finger, this scent can be readily smelled on the fingers. 
It is used to attract or excite the females. 

The three most abundant whites in the eastern and northern states are 
Pontia protodice, P. napi, and P. rapcz, the larvae of all three species being 
voracious cabbage-eaters. P. rapce, the European cabbage-butterfly, is a 
European butterfly which got to Quebec about i860 and since then has 
spread over the whole country and is the most serious pest among all the 
butterflies; it expands from if inches (male) to nearly 2 inches (female), 
has faintly yellowish-white wings with the base and apex of fore wings 
blackish and with two circular black dots on fore wings of the female and 
one in the male; there is a single black spot (in male very faint) on front 
margin of hind wings; under sides of hind wings and tip of fore wings lemon- 
yellow. P. protodice, the southern cabbage-butterfly, or checkered-white, 
has at least three black spots besides a blackish apical border on the fore- 
wings of the male, while both the wings of the female are much checkered 
with blackish brown ; the under side of the hind wings is white in the male. 
P. napi, the northern cabbage-butterfly, or mustard-white, appears in eleven 
or twelve appreciably different patterns, but characterized through all this 
variety by the pale or distinct grayish bordering of the veins; there is but 
little blackish on the wings of the male, at most one or two circular spots 
and a blackish apical border. In the western states the species of Pontia 
which will be found by most collectors are beckeri, distinguished by green 
markings on the under side of the hind wings; occidentalis, much like pro- 
todice, and sisymbri, a small species with the veins of the hind wings widely 
bordered with blackish brown on the under side. A beautiful Pierid is 
the pine-white, Neophasia menapia, of the Pacific states and Colorado; in 
both male and female the black color above is limited to the fore wings; 
there is a border along the costal margin from base to beyond the middle, 
where it bends in along the outer margin of the discal cell as a swollen club- 
like blotch; in addition the apex is broadly bordered with black in which 

446 The Moths and Butterflies 

three or four white spots appear; in some specimens the hind wings have 
a narrow broken border of scarlet on the under side. 

Of the yellows, or sulphurs, the most familiar in the eastern states is 
Eurymus philodice, the clouded sulphur, expanding i^ to 2 inches; the 
wings are pale sulphur-yellow with black outer borders and with a discal 
black spot on each fore wing and orange spot on each hind wing; in the 
female the black border of the fore wings is very broad and contains five or 
six irregular yellow spots. Similar in pattern, but with the ground color of the 
wings bright orange instead of pale yellow, is the orange- sulphur, E. eury- 
theme, common through all the West. Both of these species are polychro- 
matic and polymorphic, that is, show marked variation in ground color and 
in size, some individuals called albinos being white, some called negros 
being suffused with blackish; some are very small, others unusually large. 
A variety of names has been given to some of these aberrations because 
of their regular appearance under certain seasonal conditions. The longi- 
tudinally striped green larvae of both species feed on clover. Another com- 
mon sulphur in the southern and western states is the dog-face, large with 
pointed-tipped front wings and the yellow color of these wings so outlined 
by the black base and broad border as to produce a rough likeness to a dog's 
head seen in profile; a small discal black spot serves as the eye. The south- 
ern species is Zerene cmsonia (PI. V, Fig. 10), the Pacific coast species Z. eiiry- 
dice. The caterpillars, which are green with a whitish longitudinal stripe and 
a transverse dark line on each segment, feed on various Leguminosae. Another 
common southern and western species is Terias nicippe, the black-bordered 
orange (PL XI, Fig. 2), whose larvae feed on cassia. A striking species 
is the cloudless sulphur, Catopsila euhule, the largest of the Pierids, expand- 
ing 2^ inches; it occurs in the southern and southwestern states, its larva 
feeding on cassia. At the other extreme in size is the dainty sulphur, 
Nathalis iole, (PI. V, Fig. 7), the smallest member of the family, expanding 
but I inch ; it has the same range and food habits as the cloudless sulphur. 

In the western states occur seven or eight species of the pretty little 
Pierids known as orange-tips; only one species, Synchloe genutia (PI. XI, 
Fig. 3), is found in the east. All are small and most of them are readily 
distinguished by the characteristic orange-colored apex of the fore wings 
as shown in the colored figure of genutia. S. sara, with two named varie- 
ties, reakirtii and sara, is the commonest western species. The larvae of 
the orange-tips, so far as known, feed on Cruciferae. 

Perhaps the most striking and admired of all familiar insects are the 
great swallowtail butterflies. They have an easy, half-fluttering, half-soar- 
ing flight; their unusual size and their black and yellow (or greenish-white) 
tiger-like markings make them so conspicuous that they are fascinatingly 
apparent to the most casual observers. Twenty-one different swallowtail 

The Moths and Butterflies 


butterflies are found in the United States. Combined with them in the 
family Papilionidte are two species of curious thinly scaled black- and red- 
spotted white butterflies called parnassians, which live exclusively in high 

Fig. 639. — Swallow-tailed butterflies, Papilio rutulus. (From life; one-half 

natural size.) 

altitudes in the Rocky and Sierra Nevada Mountains. Two more species 
are found in high latitudes on this continent, namely in Alaska. Parnas- 
siiis smintheus (PI. V, Fig. 8; also Fig. 633) with four varieties is found 
in both the Colorado Rockies and Sierra Nevada, while P. clodius, a larger 


The Moths and Butterflies 

species with more translucent fore wings, is found only on the Pacific coast 
and in the Wyoming mountains. I have seen P. smintheus in great numbers 
in the beautiful fiower-dotted glacial parks of Colorado from an altitude 
of 6000 feet upward. The wings are so thinly scaled that they are nearly 
translucent, and the scales themselves are narrow and club-like, so different 
indeed from those of other butterflies that they probably have some special 
function not yet understood. The larvae are "flattened," having a some- 
what leech-like appearance; they are black or dark brown in color, marked 
with numerous light spots. The chrysalis is short and rounded at the head, 
and pupation takes place on the surface of the ground, among leaves and 
rubbish, a few loose threads of silk being spun about the spot in which trans- 
formation occurs. 

The swallowtails (Fig. 639), all except five of which belong to the genus 
Papilio (a name given them a century and a half ago by Linnaeus, the first great 

classifier of animals and plants), are readily 
distinguished by the longer or shorter "tails," 
one to three, which project backward from 
the hind wings. The ground color is black, 
sometimes suffused with metallic bluish or 
greenish, and the markings consist of yellow 
or greenish -white bands and blotches together 
with a few red, orange, and blue eye-spots on 
the upper and under sides of the hind wings. 
The larvae are large, cylindrical, fleshy, naked 
caterpillars usually conspicuously banded or 
spotted with green, black, yellow, orange, 
and- white. They are provided with a pair 
of fleshy and flexible colored "horns" (osmateria) which can be protruded 
from, or withdrawn into, the front thoracic segment and which give off a 
strong musky scent sufficiently disagreeable to repel many threatening 
enemies of the caterpillar. The chrysalids (Fig. 640) are naked, sus- 
pended by the tail from a silken button and supported by a silken girdle 
or "bridle." They often mimic very closely the coloration and surface 
configuration of the tree-trunk or other object to which they are attached 
(Fig. 640). Poulton, an English naturalist, has been able to obtain chrys- 
alids of a single swallowtail species of many different colors by enclosing 
the larvae just before pupation in separate boxes lined with paper of different 
colors. The color-tone of the chrysalid tended strongly toward that of the 
environing paper. Such a color plasticity is certainly of much advantage 
to the insect in rendering the exposed and defenceless chrysalid indistin- 
guishable. (See Chapter XVII for a discussion of "color and its uses.") 
One of the best-known butterflies of the east is the zebra swallowtail, 

v'4- n^^ 

'%«..j^ '#"^ 



Fio 640. — Chrvbdhcl of a swal- 
low-tailed buiierfly, Papilio sp 
(Natural size.) 



1 = Cercyonis alope. 

2 = Vanessa atalanta. 

3 = PapiIio cresphontes. 
4=Heodes hypophloeas. 
5 = Erynni3 sassacus. 
6=Basilarchia arthemis. 
7 = Euvanessa antiopa. 


V Mary Weilntnn, del. 

The Moths and Buttertiies 449 

Iphidides ajax (PL V, Fig. 2), which is distinguished from all other 
swallowtails by its black and greenish-white wings and its long tails; it 
appears in three forms, one, marcellus, emerging in early spring with tails 
f inch long and tipped with white; another, telamonides, appearing in 
late spring, a little larger, with tails -f inch long and bordered with white 
on each side for half the length or more, and the third the typical ajax, still 
larger, appearing in late summer and autumn. Both of the first two forms 
may come from a single brood, some of the hibernating chrysalids producing 
butterflies earlier than others. It seems to depend wholly on the time of 
issuance and not at all on the character of the parent whether an individual 
shall be of the marcellus or of the telamonides form. The ajax individuals 
are those that are produced from eggs laid in the spring by either marcellus 
or telamonides individuals. Also some few chrysalids in every brood delay 
disclosing butterflies until the next spring. " Marcellus and telamonides thus 
produce ajax the same season, or either marcellus or telamonides in the follow- 
ing spring; ajax produces itself the same season or one of the others in the 
spring; but neither marcellus nor telamonides is produced the same season 
by any of the forms" (Scudder). The larvae of this species are pea-green, 
naked, thickest in the thorax, with transverse markings consisting of black 
dots and lines and slender yellow stripes besides a yellow-edged, broad, vel- 
vety b^ack stripe on the thorax. They feed on papaw. 

Papilio turnus, the tiger swallowtail, or Turnus butterfly (PI. V, Fig. 6), 
is another common species, with a striking "negro" form called glaucus. 
In glaucus the disk of the wing is wholly dusted over with black scales so 
that the bands can be hardly seen. It is found only in regions where there 
are two or more broods a year, and is represented by females alone. The 
tiger swallowtail ranges clear across the continent, and sometimes occurs 
in great numbers; Scudder says that on a cluster of Hlacs 6g specimens were 
captured at one time by closing the two hands over them. The larvae, which 
feed on many plants but particularly like wild-cherry, are naked and leaf- 
green, with the front part of the body much enlarged and bearing a double 
stripe of yellow and black across the back, as well as a pair of yellow-black 
and turquoise eye- spots in front of this band and several rows of turquoise 
dots behind it. On the Pacific coast occur P. riitulus (Fig. 639) and P. 
eurymedon of the same general pattern of turnus, the first being black 
and yellow as turnus is, but the second being black and pale greenish or 
yellowish white. In the Rocky Mountains is found the splendid Daunus 
swallowtail, P. daunus, larger than Turnus and with two tails on the hind 
wings and a third tail-like lobe at the inner angle. The larva of rutulus 
feeds on alder and willow, of eurymedon on Rhamnus and other plants, 
and of daunus mostly on rosaceous plants. 

Of different pattern is the fine giant swallowtail, P. cresphontes (PL X, 

450 The Moths and Butterflies 

Fig. 3), native in the south, but now gradually spreading north. The 
caterpillar, sometimes called "orange-puppy" in Florida, feeds on orange- 
and lemon-trees, besides other plants, and is swollen in front of the middle, 
with the anterior part of the body rusty brown with lateral stripe, the hinder 
end of which, including two or three segments and a broad saddle in the 
middle, is cream-yellow flecked with brown. 

A smaller widely distributed and well-known PapiHo is the common 
Eastern black swallowtail, P. polyxenes, represented by five named varie- 
ties besides the type form. The black wings are crossed by two rows of 
yellow spots, the inner ones the larger, and there is a series of yellow mar- 
ginal lunules; incomplete bluish spots lie between the two yellow rows of 
spots on the hind wings, specially distinct and large in the female. The 
larva feeds on parsnips, caraway, etc., and is green-ringed with black and 
spotted with yellow. P. troiliis, the spice-bush swallowtail of the eastern 
and middle states, has a single row of well-separated yellow spots near the 
outer margin of each wing, with indications of a bluish or greenish row inside 
this, specially distinct on the hind wings; there is an orange spot at each 
end of this row on the hind wings. The larva lives on spicewood and sassa- 
fras and makes a protecting nest by tying the edges of a leaf together. The 
pipe-vine swallowtail, Laertias philenor, has no band of yellow spots, but only 
a few indicated lilac-colored remnants of spots, and has the hind wings suf- 
fused with beautiful glossy blue-green, especially beyond the base; its cater- 
pillar feeds on Dutchmen's pipe and a wild species of Aristolochia, common 
in the Appalachian forests. There are two Papilionids without tails, viz., 
Ithohahis acaiida, found in New Mexico, and I. polydamas, found in 
Florida; both are beautiful butterflies, much like P. philenor in color and 

The largest family of Rhopalocera is that of the Nymphalidse, or brush- 
footed butterflies, the vernacular name partly describing their most dis- 
tinctive structural peculiarity, namely the marked reduction (atrophy) of 
the fore legs to be functionless little hairy brush-Uke processes without tar- 
sal claws on the feet; in both sexes these fore feet lie folded on the thorax, 
"like a tippet," as Comstock has said. This and the possession of an always 
five-branched radial vein in the fore wing are about the only structural 
characteristics common to all the butterflies of this large family. The species 
range from small to large, present a bewildering variety of coloring and pattern 
and an equal variety of larval habit and appearance. All the chrysalids 
are naked, usually angular, and are suspended head downward by the tail 
without other support. Nearly 250 species of Nymphalids are recorded 
from this country, and the majority of the best-known and most abundant 
butterflies in any locality belong to the group. Some systematists consider 
the brush-footed butterflies to form several distinct families — this is the 

The Moths and Butterflies 


point of view taken by the author of the latest catalogue of North American 
Lepidoptera — while those who believe in the family unity of the group sub- 
divide it into a number of subfamilies. 

In the face of the large number of beautiful, interesting, and familiar 
species of Nymphalidae we can only select, for description in our limited 
space, a few of the most familiar and interesting. The special collector 
and student of butterflies will find awaiting him a large literature mostly 
readily available, and to this he must refer for anything like a comprehensive 
account of the species of this family. 

The all-conquering American butterfly is the monarch, Anosia plexip- 
pus (PI. XI, Fig. 4; also Fig. 641), sometimes called the milkweed-butter- 

FlG. 641. — The monarch butterfly, Anosia plexippus (above), distasteful to birds, and 
the viceroy, Basilarchia archippus (below), which mimics it. (Three-fourths natural 

fly because of the food-plant of its larva. This great red-brown butterfly 
king ranges over all of North and South America, and has begun its invasion 
of other countries by getting a foothold on the west coast of Europe and 
in almost all of the Pacific islands and in Australia. I have found the mon- 
arch the most abundant butterfly through all of the Hawaiian Islands 2000 
miles distant from the Californian coast, and still 2000 miles farther into the 
great Pacific in the Samoan Islands it is also the dominant butterfly species. 
Its success is due to its hardiness, its strong flight power, the abundance and 


The Moths and Butterflies 

cosmopolitan distribution of its food-plant, and finally and most important 
its inedibility — to birds. It secretes in its body an ill-tasting acrid fluid, 
and birds soon learn to let these disagreeable butterfly morsels alone. For 
the sake of this immunity another butterfly species, the viceroy, Basilarchia 
archippus (PL XI, Fig. i; also Fig. 641), which is not ill-tasting, mimics in 
extraordinary degree the color pattern of the monarch, so that it must be 
constantly mistaken for the disagreeable monarch and is passed unmolested 
by experienced birds. The monarch in the eastern states has a migratory 
habit not unHke that of birds, great swarms flying south in the autumn to the 
Gulf states and West Indies, returning north again in the spring, not in swarms, 
however, but singly. It ranges as far north as Canada. It has, too, a curious 
habit of assembling in great numbers in a few trees, like blackbirds or crows 
in a "roost," and hanging there quietly in masses and festoons, many indi- 
viduals clinging only to each other and not to the branches at all. On cer- 
tain great pine trees near the Bay of Monterey on the Californian coast I 
have seen myriads of monarchs thus "sembled." The eggs are laid singly 
on the leaves of various milkweed species, Asclepias cornuli the favored 
kind, and hatch in about four days. The larva (Fig. 791) attains its full 
growth in two or three weeks and is a conspicuous object with its greenish- 
white body regularly banded with narrow black and yellow stripes; it has 
two pairs of slender black filaments, one on the second thoracic and the other 
on the eighth abdominal segment. The beautiful plump chrysalid is pea- 
green, smooth, and rounded with a few black and gilt spots and bands. The 
pupal stage lasts from nine to fifteen days. There is but one generation a 
year in the north, but two appear in the south. The winter is passed by 
the adult butterfly in the warm region of the subtropics. 

Although the viceroy, Basilarchia archippus, closely resembles the 
monarch in its red-brown ground color, black-bordered veins, and small 
white spots, only one of the half-dozen other species of the same genus is 
at all like it. This one is B. floridensis found in the southern states. The 
others have a blackish ground-color with the hind wings suffused with 
greenish blue and a few conspicuous reddish blotches on the under side 
of both wings, as in the red-spotted purple, B. astyanax, common in the East, 
or broadly banded with white, as in the banded purple, B. arthemis (PI. X, 
Fig. 6), of the northeastern states, or have a blackish-brown ground with 
broad white band and red-brown apex of the fore wings, as in Lorquins 
Admiral, B. lorquini, of the Pacific states. The larvae of Basilarchia 
feed on oaks, birches, willows, currants, and various other trees and shrubs, 
and are odd-appearing caterpillars with numerous prominent tubercles or 
bosses on the back. 

Beautiful and abundant Nymphalids are the angle-wings, tawny above 
with black markings, dead-leaf-like below and often with a little silvery 

The Moths and Butterflies 


comma-spot. The comma-butterfly, Polygonia comma (PI. XI, Fig. 6; 
also Fig. 642), is a familiar eastern representative of the angle-wings. On the 
under side of each hind wing is a small but distinct silver comma or C spot. 

Fig. 642. — The comma-butterfly, Polygonia comma; two butterflies, a caterpillar, and 
empty chrysalid on gooseberry branch. (After Lugger; natural size.) 

The spiny greenish-brown larva:; feed on hops, nettles, and elms. The pale 
wood-brown chrysalids with metallic golden or silver spots are commonly 


The Moths and Butterflies 

known as hop-merchants. If the spots are golden, hops are to bring high 
prices; if silvery, low prices! The violet-tip, P. interrogationis, is another 
common eastern angle-wing and has on the under side of the hind wings a 
double silver spot a little like a question-mark but more like a semicolon. 



x7..:, ami 

' i-S. 



l^^ft^ ^^9 



V ■^ 



Fig. 643. — The larva of the violet-tipped butterfly, Polygonia interrogationis, making its 
last moult, i.e., pupating. (Photograph from Hfe by author; slightly enlarged.) 

Its chestnut-colored, pale-spotted, spiny larva feeds on hops, elms, and 
linden. Fig. 643 shows a caterpillar just pupating, and Fig. 644 shows 
the formed chrysalid. There are eight other species of Polygonia in the 
United States. 

The Vanessas are among the best known of our butterflies. Three 
species, V. atalanta (PI. X, Fig. 2), the red admiral, V. huntera, the painted 
beauty, and V. cardiii, the thistle-butterfly, are found all over the United 
States, and in addition a fourth, V . carycc, the west-coast lady, occurs on the 
Pacific coast. The latter three species are but little like atalanta, having 
the wings blackish brown, plentifully and irregularly marked with orange 
and whitish; underneath there are true eye-spots; huntera may be dis- 
tinguished from cardni by having but two complete eye-spots instead of 
several, and caryce differs from cardui by the absence of the rosy tint peculiar 
to that species, the tawnier ground-color of the upper surfaces, and the com- 
plete black band which crosses the discal cell of the fore wings. Atalanta 



1 = Basilarchia archippus. 

2 = Terias nicippe. 
3=Synchloe genutia. 
4=Anosia plexippus. 
5 = Anasa andria. 
6=Polygonia comma. 


Mary IVeUman, del. 

The Moths and Butterflies 


and cardiii occur also in Europe, and cardui is held to be the most nearly 
cosmopolitan of all butterflies, ranging over nearly the whole earth outside 
the arctic and antarctic regions. Its larvai feed on thistles by preference, 
but on almost any composite if necessary: those of huntera on everlasting 
and other Gnaphalieas; those of atalanta on nettles; vi^hile those of caryce 
feed on Lavatera assiirgenliflora. All these larvae are spiny. 

Tvi^o striking, widely distributed, and abundant butterflies are the mourn- 
ing-cloak, Euvanessa aniiopa (PI. X, Fig. 7), and the peacock-butterfly, 
or buckeye, Junonia cxnia (PI. V, Fig. i). Both are found over nearly 
all of our country, and the mourning-cloak is common in Europe. The 

Fig. 644. — Chrysalid or pupa of the violet-tipped butterfly, Polygonia intcrros^ationis. 
(Photograph from life by author; sUghtly enlarged.) 

larva of the buckeye is black-gray marked with minute black-edged orange 
dashes and dots transversely arranged, and has long spines all over its body; 
it feeds on Scrophulariaceae, especially Gerardia. The larva of the mourning- 
cloak is velvety black sprinkled with white papilla? and with a row of large 
medio-dorsal orange spots, and has spines much longer than the body seg- 
ments. A curious butterfly of the Mississippi Valley and Great Plains 
is An(^a andria, the goatweed-butterfly (PI. XI, Fig. 5). The larva, 
which is naked, gray-green, and studded with numerous paler points, feeds 
on species of Croton, the goatweeds. The American tortoise-shell, Aglais 

456 The Moths and Butterflies 

milberti, which occurs commonly in the North, has brownish-black wings 
with a broad orange fulvous band between the middle and outer margin; 
there are also two fulvous spots in the discal cell of the fore wing. The 
larva, which feeds on nettles, is spiny, velvety black above, greenish yellow 
below, and profusely dotted with whitish spots or points. Another northern 
butterfly is the Compton tortoise, Eiigonm j-album, which resembles in 
general color and pattern the angle-wings (Polygonia), but has the hinder 
margin of the fore wings straight, the markings on these wings heavier, 
and a whitish spot on both fore and hind wings near the apex; there is also a 
small L-shaped silver spot on the under side of the hind wings. Eugonia 
calijornica, the California sister, is a beautiful butterfly common on the 
Pacific coast and found occasionally in the Rocky Mountains; it is velvety 
blackish brown with a broad white transverse bar across each wing, inter- 
rupted on the fore wings and tapering out on the hind wings, and with a 
conspicuous large orange-brown patch nearly filling the apex of the fore 
wings. Its larva feeds on oaks. 

Two large groups of brush-footed butterflies, some of whose species 
occur in every locahty, are the fritillaries, or silver-spots (genus Argynnis 
and allies) and the checker-spots (genus Melitaea and allies). The 
fritillaries, mostly medium-sized to large butterflies, are usually red-brown 
with numerous black spots scattered over the upper surface of both wings; 
the hind wings usually bear on the under side a number of striking silvery 
blotches, which give these butterflies their name of silver-spots. The regal 
fritillary, Speyeria idalia, of the Atlantic states, expands 2f to 4 inches and 
has the fore wings bright fulvous above spotted with black, and the hind 
wings blue-black with a marginal row of fulvous and submarginal row 
of cream-colored spots; both fore and hind wings have silver blotches on 
the under sides. The black, ocher, and red-banded caterpillars have six 
rows of fleshy black and white spines; they feed on violets and are nocturnal. 
The spangled fritillary, Argynnis cyhele, is a good example of the more 
usual coloring and pattern of the group. It expands from 3 to 4 inches, 
has both wings fulvous above and thickly spotted with black; the under 
side of the hind wings is silver-blotched; in the female the basal half of 
the fore and hind wings above is dark chocolate-brown. The caterpillar 
is black with six rows of shining black branching spines, and feeds on violets. 
Numerous other smaller Argynnids are like cybele in color and pattern: 
it is difficult to distinguish the various species. 

The checker-spots, small to medium size, blackish with red and yellowish 
spots, are represented by numerous species in the western mountain states, 
but by only two species in the east. The Baltimore, Euphydryas phaeton, 
expanding if to 2^ inches, is the most familiar eastern checker-spot; it is 
black above with a marginal row of red spots followed by three rows of pale- 

The Moths and Butterflies 457 

yellow spots on the fore wings and two on the hind wings; besides there 
are some scattered red spots and some other yellow ones. The caterpillar 
is black, spiny, and banded with orange-red; it feeds chiefly on Chelone 
glabera, a kind of snakehcad. On the Pacific coast the chalcedon, 
Melitaea chalcedon, is the most abundant checker-spot, although several 
other species are common. It has black wings spotted with red and 
ocher-yellow; the spiny black caterpillar feeds chiefly on Mimulus and 

The satyrs or meadow-browns are a group of fifty or more beautiful velvet- 
brown butterflies whose markings consist chiefly of eye-spots, large and small, 
on both upper and under wing surfaces. A number of species are abundant 
and familiar, but a majority live exclusively in mountain states, and especially 
in the west. The common wood-nymph, or eyed grayling, Cercyonis dope, 
(PI. X, Fig. i), is the most familiar eastern and middle state species. 
A larger similarly patterned form, C. pegala, is common in the south. The 
larvae of the meadow-browns feed on grasses, are pale green or light brown, 
and have the last abdominal segment forked. On the Pacific coast one 
of the most abundant autumn butterflies is the California ringlet, Cceno- 
nympha calijornica, a small buffy-white member of this group with small 
eye-spots only on the under side of the wings. A number of interesting 
butterflies related to the meadow-browns are found only on mountain-tops 
or in high latitudes (arctic region) the equivalent in Hfe conditions of high 
altitudes. In the Rocky Mountains on the peaks of the Front Range (13,000 
feet altitude) I have struggled, gasping in the thin air, after beautiful frail 
little brown and grayish butterflies, CEneis and Erebia. Far above timber- 
line on bleak mountain-tops, masses of broken granite overspread for great 
spaces with lasting snow, these hardy little flutterers live successfully. At 
the edges of the great snow-fields are patches of alpine flowers, fragrant 
dwarf forget-me-nots and buttercups, which furnish food and interest for 
them in the solitude of the high peaks. 

The mountain-top butterflies of the White Mountains, of the Rocky 
Mountains, and of the Sierra Nevada are closely allied; indeed individuals 
of the same species are found on the summit of Mt. Washington and on 
the crest of the Rockies, and nowhere between these two widely separated 
localities. The question as to how this interesting condition of things came 
about would be answered (by the student of distribution) as follows: In 
glacial times the species probably ranged clear across the continent. With 
the retreat of the great continental ice-sheet, while most of the butterflies 
followed it closely north, or became in successive generations slowly adapted 
to the temperate life conditions, some few probably followed up the slowly 
retreating local mountain glaciers. In time, therefore, the descendants 
of these arctic-loving species found themselves still under truly arctic con- 


The Moths and Butterflies 

ditions on the snow-covered mountain-tops, but isolated by the temperate 
lowlands from the rest of their kind on other mountain-tops or in arctic 

There are several excellent books about American butterflies which will 
help the nature student classify his specimens, and tell him of the distribution 
and habits of the various species. Among the best are Comstock's "How 
to Know the Butterflies," Holland's "The Butterfly Book," and Scudder's. 
' ' Everyday Butterflies." 




AND ANTS (Order Hymenoptera) 

EES, ants, and wasps are the familiar Hymenoptera. 
They are the "intelhgent" and the "social" in- 
sects, and therefore seem, of all the insect hosts,, 
those living the most speciaHzed or "highest" kind 
of life. As intelligence and social life are precisely 
those characteristics of our own which most dis- 
tinctly set us off from other animals, we are quick 
to appreciate the worth of similar attributes in the 
"ant and bee people." But in actual degree of 
specialization of instinct 
and behavior the perform- 
ances of the soHtary wasps 
and bees are little less wonderful than those of 
the social kinds, and the amazing character of the 
life-history of many of the obscure and unfamiliar 
parasitic and gall-making Hymenoptera ought to 
incite as much interest and scientific curiosity as the 
marvels of the bee community. The Hymenoptera 
constitute a large order, 7500 species in this coun- 
try, and one of endless variety of habit and struc- 
ture. Few generalizations indeed can be made that 
will apply to all the members of the order, although 
there is no question concerning the true relationship 

of all the kinds of insects included in the order. Of ^^^- 645 --Mouth-parts of a 

honey-bee with maxilla 
the structural characteristics common to the Hymen- and mandible of right side 

optera the clear, membranous condition of the two removed, w J., mandible; 

. . . mx., maxula.; mx.p., max- 

pairs of wings gives the name to the order {hymen, juary palpus; mx.L, max- 

membrane; pteron, wing). The front wings are iHary lobe; st., stipes of 
1 ^1 ^1 1 • 1 1 11 • I J -^1 maxilla; cJ., cardoof max- 
larger than the hind ones, and all are provided with jjj^. //. labium- sm. sub- 
comparatively few branched veins, whose homologies mentum of labium; m., 

have not been fully worked out. The workers "^^ntum of labium; pg., 
•^ . . paraglossa; gl., glossa; 

(infertile females) of all the ant species are wingless, H.p., labial palpus. 



Saw-flies, Gall-flies, Ichneumons, 

.as are also the females of the Mutillid wasps and a few other exceptional 

forms. In many Hymenoptera (shown 

well in the honey-bee) the fore 
(costal) margin of the hind wings 
bears a series of small but strong 
recurved hooks which, when the 
wings are outspread, fit snugly over 
a ridge along the hind margin of the 
fore wing, the two wings of each side 
being thus fastened together so as to 
move synchronously. A structural 
characteristic not readily made out 
but of much morphological impor- 
tance is the complete fusion of the 
Fig. 646.— Lateral aspect of head of full- true first abdominal segment with 
grown larva of honey-bee which has been ^j^^ thoracic mass, SO that the small 
cleared so as to show the forming adult head ... , , 

within, ih., head of aduU; i.e., compound articulating segment between what 
eye of adult; Ic, body-wall of larval head; ^^.g called thorax and abdomen if 
i ant., antenna of adult;, mandible of ,, ,, j i_ j • 1 

larva;, mandible of adult;, really the second abdominal seg- 
maxilla of larva;, maxilla of adult; nient., labium of larva;, labium of adult. 

The mouth-parts are variously 
modified, but usually are fitted for both biting and sucking (or lapping). 
This is arranged for by having the maxilla^ and 
labium more or less elongate and forming a sort 
of proboscis for taking up liquids, while the man- 
dibles always retain their short, strong, toothed, 
jaw-like character. The mandibles of the honey- 
bee are modified into admirable little "trowels" 
for moulding wax and propolis. The females 
throughout the order are provided either with a 
saw-like or boring or pricking ovipositor, or with 
the same parts modified to be a sting. The sting 
is possessed by the wasps, bees, and ants (rudi- 
mentary in many ants), on which account these 
groups are often referred to collectively as the 
aculeate Hymenoptera. The sting of the honey- 
bee is shown in Fig. 650 and is a well-developed 
example of this characteristic hymenopterous 
weapon of defence and offence. The barb-tipped 
darts (d) extend down through the sheath (s) and 
are controlled by the chitinous bars called levers 
(/). The poison produced in the poison-gland ( and stored in the 

Fig. 647. — Mouth-parts of 
mud-wasp, with mandible 
and maxilla of right side 
removed, md., mandible; 
mx., maxilla; mx.l., max- 
illary lobe; inx.p., maxil- 
lary palpus; //., labium; 
in., mentum of labium; 
pg., paraglossa; gl., glossa; 
li.p., labial palpus. 


1 = Spharophthalraus californicus. 

2 = Polistes aurifer. 
3=Stizus sp. 
4=Psithyrus elatus. 
5 = Bomb us vagans. 
6=Agapostemon radiata. 
7 = Xylocopa virginica. 
8=Bembex spinolae. 
9=Vespa germanica. 

io= Bombus californicus. 
11 = Anthophora pacifica. 
i2 = Polybia flavitarsis. 
i3=Chalybion ccjeruleura. 
i4=Sphex ichneunionea. 
15 = Pelopeus servilla- 

Ma^V Wellni,<n. del. 

Wasps, Bees, and Ants 


sac (p.s.) flows from this into lesser reservoirs in the expanded base of the 
sheath and escapes through the valve (v) along the darts 
into the wound. The tactile (and perhaps olfactory) palpi 
(p) are used to explore the surface of the object to be 
stung. The modifications of the various appendage-like 
parts which compose the sting to form an egg-depositing 
organ (ovipositor) are extremely various and are described 
later in connection with various special groups. The 
number of separate parts or processes which compose 
the ovipositor or sting and which arise from the two ab- 
dominal segments next in front of the terminal one is 
six, and some entomologists consider these parts to be true 
appendages, homologous with the legs and mouth-parts. 
In the development of all Hymenoptera the meta- 
morphosis is complete, and the larva? are, more than 
in any other order, helpless and dependent for their 
food and safety on the provision or care of the parents 

"^ \mr.l. 
Fig. 64S. — Frontal as- 
pect of head of larva 
of mud-wasp, md., 
mandible; mx., max- 
illa; mx.l., maxillary 
lobe; //., labium; 
li.p., labial palpus. 

With many 


Fig. 649. Fig. 650. 

649. — Lateral aspect of head of full-grown larva of mud-wasp cleared so as to 
show forming adult head within, i.h., head of adult; i.e., compound eye of adult; 
I.e., body-wall of larval head; iant., antennae of adult;, mandible of larva;, mandible of adult;, maxilla of larva;, maxilla of adult;, 
maxillary palpus of adult; I. It., labium of larva;, labium of adult; li.lip., labial 
palpus of adult. 
Fig. 650. — Sting of the worker honey-1 ee. p-gl., poison-gland; p.s., poison-sac; d., dart; 
/., levers; v., valve; s., sheath; p., palpus. 

species, as the solitary wasps and bees, food is stored up in the cell in which. 

462 Saw-flies, Gall-flies, Ichneumons, 

the egg is deposited, so that the larva on hatching will find it ready to hand. 
With the social wasps and bees and all the ants, the workers bring food to 
the larva during its whole life. With the lower forms, the parasitic and 
gall-making kinds, the egg is deposited on or in a special and suflScient food- 
supply. All these unusual conditions are described in the discussion of 
the various groups. Indeed this whole chapter on the Hymenoptera is writ- 
ten especially with the aim of illustrating the biology, the special life con- 
ditions and relations of the various larger groups of these insects, rather 
than with the aim which determined the character of the chapters on the 
beetles (Coleoptera) and moths and butterflies (Lepidoptera), namely, that 
of presenting a systematic survey of the classification and individual habits 
of those members of the order most likely to be seen or captured by the col- 
lector. The beetles and the moths and the butterflies are the insects which 
fill the cabinets of the amateur and beginning student, and names and facts 
concerning particular species are likely to be the particular desiderata in 
connection with them. But it is the extraordinary and "wonderful" char- 
acter of the ecological relations and physiological adaptations of the Hymen- 
optera which make these insects of such interest to nature-lovers, and which, 
indeed, is the subject that can most profitably be given special attention 
by any student of the order. Without, therefore, making any further attempt 
to formulate generalizations concerning this great complex of variously 
mannered insects, we may begin our study of its members arranged in sub- 
ordinate groups, this grouping depending rather upon general biologic char- 
acteristics than strictly classific ones. 

The classification of the Hymenoptera is a matter that interests but few 
amateurs; only a few families are at all well represented in general collec- 
tions. Distinction among the more familiar larger groups, as the ants, bees, 
wasps, saw-flies, horn-tails, and ichneumons, is usually pretty well marked 
in the general habitus or tout ensemble of appearance. Certain other of the 
larger groups, composed of minute parasitic species, are almost unknown 
to the general collector; indeed but two or three American professional 
entomologists would attempt to distinguish species in these groups. In the 
following table, therefore, and in the later discussion of the various groups, 
I have lumped these little-known families together on a basis of common- 
ness of habit, namely, of parasitic life, and devoted the space to a general 
account of the extraordinary life-history and habits which these parasitic 
Hymenoptera have adopted, with some reference to the special habits of 
certain particular species. Their classification into smaller groups is left 

Wasps, Bees, and Ants 463 


A. Trochanters (segment between the rounded basal coxa and the long femur) of 
the hind legs divided in two, i.e., two-segmented; female with a saw or borer at 
tip of body for depositing the eggs. 
B. Abdomen joined broadly to the thorax. 

C. Tibiae of fore legs with two apical spurs; female with a pair of saw-like 
egg-depositing processes at tip of abdomen. 

(Saw-flies.) Family Tenthredinid^ (p. 464). 
CC. Tibias of fore legs with one apical spur; female with elongate borer 

instead of saw (Horn-tails.) Family SlRlciD^ (p. 466). 

BB. Base of abdomen constricted, so that it joins the thorax as if by a stem. 
C. Abdomen joined to the dorsum of the metathorax. 

(Ensign-flies.) Family Evaniid^. 
CC. Abdomen joined to posterior aspect of metathorax. 

D. Fore wings with few veins and no closed cells (a few exceptions); 
very small parasitic Hymenoptera. 

Families Chalcidid^ and Proctotrypid^ (p. 476). 
DD. Fore wings with one or more closed cells (a few exceptions). 
E. Fore wings without a stigma (Fig. 655). 

(Gall-flies.) Family Cynipid^ (p. 467). 
EE. Fore wings with a stigma (Fig. 671); parasitic Hymenop- 
tera, from very small to large. 

(The Ichneumons and other parasites.) Families Braco- 
NiD^, Stephanid.e, Ichneumonid^, and Trigonalid^ (p. 476). 
AA. Trochanters of hind legs not divided, i.e., consisting of a single segment; female 
often with a sting. 
B. Fore wings with no closed submarginal cells (Fig. 683). 

C. Abdomen long and slender, and antennae also long and filiform. 

Family Pelecinid^ (p. 484). 
CC. Abdomen short, but little longer than head and thorax; antennae short 

and elbowed (Cuckoo-flies.) Family Chrysidid^ (p. 498). 

BB. Fore wings with at least one closed submarginal cell. 

C. First abdominal segment and sometimes the second segment in the shape of 
a small disk-like piece (Fig. 743). 

(Ants.) Superfamily Formicina (p. 533). 
CC. Basal segment (or segments) of abdomen normal or elongated to form 
a peduncle. 

D. First segment of tarsus of hind legs cylindrical and naked or with 
but little hair. 
E. Wings not folded longitudinally when at rest. 

(Digger-wasps.) Superfamily Sphecina (p. 490). 
EE. Wings folded longitudinally when at rest. 

(True wasps.) Superfamily Vespina (p. 503). 
DD. First segment of tarsus of hind legs expanded and flattened and